Seed depredation negates the benefits of midstory hardwood removal on longleaf pine seedling establishment

Midstory hardwoods are traditionally removed to restore longleaf pine on fire‐excluded savannas. However, recent evidence demonstrating midstory hardwood facilitation on longleaf pine seedling survival has brought this practice into question on xeric sites. Also, midstory hardwoods could facilitate longleaf pine seedling establishment, as hardwood litter may conceal seeds from seed predators or improve micro‐environmental conditions for seedling establishment. However, little is known about these potential mechanisms. In this study, we tracked longleaf pine seed depredation and germination in artificially seeded plots (11 seeds/m²) in a factorial design fully crossing hardwood retention or removal with vertebrate seed predator access or exclusion in the Sandhills Ecoregion of North Carolina, U.S.A. Seed depredation averaged 78% across treatments and was greatest in unexcluded plots. Hardwood retention did not affect seed depredation. Longleaf pine averaged 3.6 germinants/4 m² across treatments, and was six times more abundant where vertebrates had been excluded. Hardwood removal had a strong positive effect on seedling germination, likely due to the removal of litter, but only when vertebrates were excluded. Our results indicated midstory hardwoods are not facilitating longleaf pine seedling establishment. Nevertheless, our results indicated that hardwood removal may not increase longleaf pine seedling establishment, as seed depredation diminished the effectiveness of hardwood removal under mast seed availability. Collectively, these results demonstrate the underlying complexity of the longleaf pine ecosystem, and suggest that planting may need to be part of the restoration strategy on sites where seed depredation limits longleaf pine natural regeneration.     

Effects of Natural Barriers and Habitat on the Western Spread of Raccoon Rabies in Alabama

Abstract: Although domestic animal transmission of rabies has largely been mitigated, the disease remains a concern in both Europe and North America where wildlife transmission has caused epizootics. Raccoon (Procyon lotor) rabies was established in Alabama, USA, in 1975, primarily in the southeastern corner of the state. However, with the exception of isolated events, rabies has not continued to spread westward across the Alabama River. We monitored movements of 100 radiocollared raccoons on 2 sites within hardwood and agriculture habitats in a rabies enzootic area east of the Alabama River, in managed pine habitat area west of the river where rabies sporadically occurs, and in a mixed pine hardwood area outside of the known rabies enzootic area to determine if raccoon movements and habitat use in certain habitat types and the presence of a river may serve as natural barriers preventing the western spread of rabies in Alabama. We also examined raccoon contact rates to determine if they influence disease transmission through static and dynamic interactions. Raccoons in mixed pine-hardwood forest habitats had smaller home ranges and less overlap of ranges compared to the other 3 habitats. However, static interactions between habitats in the use of overlap areas did not differ (F_11,129 = 1.63, P = 0.09). Rabies antibody titers were highest in the managed pine habitat (28%) even prior to oral vaccine bait distributions in spring of 2004 and 2005. Biomarker data from radiocollared and additional raccoons captured after the bait distribution west of the Alabama River demonstrated a low efficacy of the vaccine reaching the small southern raccoons. The combination of the river as a partial barrier, the high percentage of pine forested habitat west of the river, and limited spatial movements of raccoons within these forested habitats appears to have reduced the likelihood of rabies establishing west of the river. Understanding different host-habitat- disease systems is important for successful management of diseases. Based on our results, we recommend that the oral vaccine program continue to use the Alabama River as a partial barrier and baiting be concentrated in the fragmented bottomland hardwood forests and around larger bodies of water where raccoon densities are highest. Success of baiting strategies designed to take advantage of northern raccoon dynamics and habitat use may not be applicable to southern populations.     

Structure and Composition of Historical Longleaf Pine Ecosystems in Mississippi,USA

Longleaf pine (Pinus palustris) historically was a widespread ecosystem composed of a simple tree canopy and grasslands ground layer. After widespread loss of this ecosystem due to logging and fire exclusion, little quantitative information exists about historical structure for restoration goals. We identified composition in De Soto National Forest and Pearl River County, Mississippi, USA, and density, basal area, and percent stocking in Pearl River County using General Land Office surveys and US Forest Service Forest Inventory and Analysis surveys. Historical longleaf ecosystems were about 85% pine, with lesser amounts of broadleaf evergreen and oak species. Densities were about 175 to 180 trees/ha, mean tree diameters were 45 cm, and stocking was around 60% to 65%, which suggested longleaf pines were closed woodlands. Current forests are 38% to 57% pine, primarily loblolly, while longleaf pine is 2% to 8% of composition. Indeed, current longleaf pine composition across the Coastal Plain averages 3% and does not reach 10% at smaller landscape scales. Fire-sensitive broadleaf species of water oak, sweetgum, yellow-poplar, and red maple increased from about 0.5% composition to 2% to 10% of composition. Forests became twice as dense, at about 280 trees/ha to 330 trees/ha, with mean tree diameters of 22 cm. These results characterize conversion from open old growth longleaf forests, resulting in part from human maintenance, to successional forests due to human disruption of the historical ecosystem. It is important to remember structure and composition of historical forests for restoration and recognize wholesale changes so that successional forests do not become the new social and cultural baseline.     

Longleaf pine (Pinus palustris Mill.) fire scars reveal new details of a frequent fire regime

Question: How frequent and variable were fire disturbances in longleaf pine ecosystems? Has the frequency and seasonality of fire events changed during the past few centuries? Location: Kisatchie National Forest, Western Gulf Coastal Plain, longleaf pine–bluestem ecosystem, in relatively rough topography adjacent to the Red River, Louisiana, USA. Methods: Cross‐sections of 19 remnant pines exhibiting 190 fire scars were collected from a 1.2‐km² area. Tree‐rings and fire scars were precisely dated and analysed for the purpose of characterizing past changes in fire and tree growth. Temporal variability in fire occurrence and seasonality was described for the pre‐ and post‐European settlement periods. Seasonality of historic fires was determined by the scar position within the rings. The relationship between fire and drought was investigated using correlation and superposed epoch analysis. Results: The mean fire return interval for the period 1650‐1905 was 2.2 years (range 0.5 to 12 yr). Significant new findings include: evidence for years of biannual burning, temporal variability in fire seasonality, an increase in fire frequency and percentage of trees scarred circa 1790, and synchronous growth suppression and subsequent release of trees coinciding with land‐use changes near the turn of the 20th century. Drought conditions appeared unrelated to the occurrence of fire events or fire seasonality. Conclusions: Multi‐century fire history records from longleaf pine ecosystems are difficult to obtain due to historic land‐use practices and the species high resistance to scarring; however, our results indicate potential for reconstructing detailed fire histories in this ecosystem. Fire scars quantitatively documented one of the most frequent fire regimes known. Fire regime information, such as the temporal variability in fire intervals, prevalence of late‐growing season fire events and biannual burning, provide a new perspective on the dynamics of longleaf pine fire regimes.     

Canopy thinning,not agricultural history,determines early responses of wild bees to longleaf pine savanna restoration

Longleaf pine savannas are highly threatened, fire‐maintained ecosystems unique to the southeastern United States. Fire suppression and conversion to agriculture have strongly affected this ecosystem, altering overstory canopies, understory plant communities, and animal populations. Tree thinning to reinstate open canopies can benefit understory plant diversity, but effects on animal communities are less well understood. Moreover, agricultural land‐use legacies can have long‐lasting impacts on plant communities, but their effects on animal communities either alone or through interactions with restoration are unclear. Resolving these impacts is important due to the conservation potential of fire‐suppressed and post‐agricultural longleaf savannas. We evaluated how historical agricultural land use and canopy thinning affect the diversity and abundance of wild bees in longleaf pine savannas. We employed a replicated, large‐scale factorial block experiment in South Carolina, where canopy thinning was applied to longleaf pine savannas that were either post‐agricultural or remnant (no agricultural history). Bees were sampled using elevated bee bowls. In the second growing season after restoration, thinned plots supported a greater bee abundance and bee community richness. Additionally, restored plots had altered wild bee community composition when compared to unthinned plots, indicating that reduction of canopy cover by the thinning treatment best predicted wild bee diversity and composition. Conversely, we found little evidence for differences between sites with or without historical agricultural land use. Some abundant Lasioglossum species were the most sensitive to habitat changes. Our results highlight how restoration practices that reduce canopy cover in fire‐suppressed savannas can have rapid benefits for wild bee communities.     

Post‐fire development of faunal habitat depends on plant regeneration traits

The concept that vegetation structure (and faunal habitat) develops predictably with time since fire has been central to understanding the relationship between fire and fauna. However, because plants regenerate after fire in different ways (e.g. resprouting from above‐ground stems vs. underground lignotubers), use of simple categories based on time since fire might not adequately represent post‐fire habitat development in all ecosystems. We tested the hypothesis that the post‐fire development of faunal habitat structure differs between ecosystems, depending on fire regeneration traits of the dominant canopy trees. We measured 12 habitat components at sites in foothill forests (n = 38), heathy woodlands (n = 38) and mallee woodlands (n = 98) in Victoria, Australia, and used generalised additive models to predict changes in each variable with time since fire. A greater percentage of faunal habitat variables responded significantly to time since fire in mallee woodlands, where fires typically are stand‐replacing, than in foothill forests and heathy woodlands, where canopy tree stems generally persist through fire. In the ecosystem with the highest proportion of epicormic resprouters (foothill forests), only ground cover and understorey vegetation responded significantly to time since fire, compared with all but one variable in the ecosystem dominated by basal resprouters (mallee woodlands). These differences between ecosystems in the post‐fire development of key habitat components suggest there may also be fundamental differences in the role of fire in shaping the distribution of fauna. If so, this challenges the way in which many fire‐prone ecosystems currently are categorised and managed, especially the level of dependence on time since fire and other temporal surrogates such as age‐classes and successional states. Where time since fire is a poor surrogate for habitat structural development, additional complexity (e.g. fire severity, topography and prior land‐use history) could better capture processes that determine faunal occurrence in fire‐prone ecosystems.     

Conservation management of Pinus palustris ecosystems from a landscape perspective

Question: Do case studies from silvicultural and restoration studies and applied conservation management in second‐growth Pinus palustris stands provide unique insights for conservation models? Methods: A review of management paradigms that conserve the high biological diversity and rare species, drawn from characteristics in both second‐growth and old‐growth stands, is presented for fire‐maintained Pinus palustris (longleaf pine) forests. Results: A common assumption that old‐growth stands provide the primary information for the development of conservation management strategies de‐emphasizes lessons learned from second‐growth and restoration studies. Primary conservation management goals for the Pinus palustris ecosystem include the perpetual regeneration of the fire‐maintained forest and conservation of the characteristically high biological diversity and rare species. Several attributes, such as a sustained population of Picoides borealis (red‐cockaded woodpecker), Aristida stricta (wiregrass)‐dominated ground cover, and undisturbed upland‐wetland ecotones, can predict a diverse and ecologically functional ecosystem. Such indicators are linked to critical structural and functional features of the system and reflect previous land management histories that suggest sustainable approaches. Conclusions: A traditional definition of ‘old‐growth’ relying on overstorey may be limited in describing important features of healthy, diverse Pinus palustris ecosystems. Some characteristics are significantly more important for maintenance of diversity than age of the trees and these features may be present in old‐ or second‐growth forests. We advocate that the management history, structural characteristics and landscape context of stands that harbour desirable conservation attributes (red‐cockaded woodpeckers, wiregrass, gopher tortoises and undisturbed upland‐wetland ecotones) can be used as indicators to identify important conservation and forest management principles.     

Vegetative Characteristics of Recently Reforested Bottomlands in the Lower Cache River Watershed, Illinois, U.S.A.

Abstract Interest in restoring native ecosystems is resulting in conversion of marginal agricultural lands to bottomland hardwood‐dominated forests in the midwestern and midsouthern United States. Growing stock for these efforts typically consists of planted oak (Quercus spp.) and volunteer vegetation. Reports of mixed reforestation success and the lack of post‐establishment tree growth data prompted this evaluation of vegetation characteristics of 5‐ to 7‐year‐old operational restorations in the Lower Cache River Watershed in southernmost Illinois, U.S.A. Fraxinus pennsylvanica (green ash), Acer negundo (box‐elder), and Liquidambar styraciflua (sweetgum) together comprised 77% of all tree stems observed. Full stocking of overstory tree species can be expected to produce a closed canopy stand within 160 m of a forested edge, due primarily to the abundance of rapidly growing volunteer‐origin trees. Planted oaks contributed minimally to total tree stocking but were present in sufficient numbers to eventually improve wildlife habitat, and therefore satisfied restoration objectives. Oak height was 23% greater when in the presence of a non‐oak tree species. Herbaceous cover was dominated by Solidago gigantea (late goldenrod) and Juncus spp. (rushes). Solidago gigantea was associated with poor growth and low density of non‐oak stems, whereas Juncus dudleyi (Dudley's rush) was associated with taller non‐oak stems. These results suggest that the presence of volunteer‐origin trees is crucial for the creation of full stand stocking that will result in rapid development of a closed canopy forest. Improved success of future reforestation efforts will require more intensive methods to establish adequate stocking beyond 160 m of a forest edge. Methods described here could be adapted for agricultural field to forest restorations in other regions to predict critical distances from volunteer seed sources within which supplemental planting would be unnecessary to meet tree stocking objectives.     

Canopy Closure and Emigration by Juvenile Gopher Frogs

ABSTRACT Although studies have addressed effects of abrupt transitions in habitat type (e.g., forest-clear-cut or forest-field edges) on amphibian movements, little is known about effects of more subtle habitat transitions on patterns of migration and habitat use in amphibians. We used radiotelemetry to study movement patterns of juvenile gopher frogs (Rana capito) emigrating from ponds that were surrounded by longleaf pine (Pinus palustris) forest that varied in structure as a result of fire suppression. Our primary purpose was to determine if frogs emigrate directionally from their natal ponds and select habitat at random during their first month following metamorphosis. We found that frogs emigrated in nonrandom directions from ponds that were surrounded by heterogeneous habitat and selected fire-maintained habitat that was associated with an open canopy, few hardwood trees, small amounts of leaf litter, and large amounts of wiregrass (Aristida beyrichiana). Fire-maintained habitat contained higher densities of burrows excavated by gopher tortoises (Gopherus polyphemus) and small mammals, which are the priamry refuge sites for both juvenile and adult gopher frogs. Frogs moved up to 691 m from their natal ponds, frequently crossed dirt roads, and even seemed to use these roads as migration corridors. To maintain suitable terrestrial habitat for gopher frogs, including habitat used by migrating individuals, it is important to apply frequent prescribed fire to uplands surrounding breeding ponds that lead all the way to the edges of breeding ponds, as well as through ponds during periodic droughts.     

Restoring Fire to Long-Unburned Pinus palustris Ecosystems: Novel Fire Effects and Consequences for Long-Unburned Ecosystems

Biologically rich savannas and woodlands dominated by Pinus palustris once dominated the southeastern U.S. landscape. With European settlement, fire suppression, and landscape fragmentation, this ecosystem has been reduced in area by 97%. Half of remnant forests are not burned with sufficient frequency, leading to declines in plant and animal species richness. For these fire‐suppressed ecosystems a major regional conservation goal has been ecological restoration, primarily through the reinitiation of historic fire regimes. Unfortunately, fire reintroduction in long‐unburned Longleaf pine stands can have novel, undesirable effects. We review case studies of Longleaf pine ecosystem restoration, highlighting novel fire behavior, patterns of tree mortality, and unintended outcomes resulting from reintroduction of fire. Many of these pineland restoration efforts have resulted in excessive overstory pine mortality (often >50%) and produced substantial quantities of noxious smoke. The most compelling mechanisms of high tree mortality after reintroduction of fire are related to smoldering combustion of surface layers of organic matter (duff) around the bases of old pines. Development of effective methods to reduce fuels and competing vegetation while encouraging native vegetation is a restoration challenge common to fire‐prone ecosystems worldwide that will require understanding of the responses of altered ecosystems to the resumption of historically natural disturbances.     

Are raccoons limited by the availability of breeding resources? Evidence of variability in fecundity at fine spatial scales

Despite their behavioral plasticity, recent research suggests raccoons (Procyon lotor) exhibit variability in demography and genetic structure among individual habitat patches in landscapes heavily affected by anthropogenic land use. Consequently, elucidation of vital rates at fine-spatial scales is needed to implement appropriate management strategies for this species. To evaluate the degree of variability in productivity that exists among individual habitat patches, we collected reproductive (n = 170) and cementum annuli age data (n = 383) for raccoons occupying 30 forest patches varying in local and landscape-level habitat attributes within a highly fragmented agricultural ecosystem. Across all females sampled, pregnancy rates averaged 85% but were highly variable among ages (range: 47–100%). Average litter sizes ranged from 3.2 to 4.7, but did not differ as a function of age. At the landscape-level, we observed significant variability among habitat patches in the total number of offspring produced (range: 0–80), indicating that individual patches vary in their contribution to the overall size and stability of the global (landscape-level) population. Variability in productivity among habitat patches primarily was driven by local differences in the availability of denning resources, likely because of the influence of this variable on variability in the number, age structure, and reproductive rate of females. Our results suggest that in agricultural ecosystems, the reproductive potential and temporal stability of raccoons within individual habitat fragments is inextricably linked to the density of tree cavities. © 2012 The Wildlife Society.     

The Lack of Favorable Responses of an Endangered Pitcher Plant to Habitat Restoration

Restoring habitat structure that existed before active and inadvertent fire suppression is thought to be critical to maintaining populations of some rare plants in fire‐suppressed habitats. Nevertheless, the impacts of habitat restoration on most endangered plants are poorly understood. Current theory predicts and empirical studies have shown that the reduction of shade or competition (frequently a goal of many habitat restoration projects in degraded fire‐dependent ecosystems) benefits plants adapted to nutrient‐poor soils by increasing the benefit‐to‐cost ratio of adaptations for enhanced nutrient capture. Here, I examined how experimental reduction of neighboring plants in a wet longleaf pine community dominated in the ground cover by shrubs and stump sprouts influenced the growth, the reproduction, the carnivorous effort, and the benefits of carnivory in a U.S. federally endangered species, Sarracenia rubra ssp. alabamensis. Two years of data showed no significant effects of neighbor reduction or prey exclusion on any of several indicators of plant performance, nor was there any evidence of a hypothesized morphological trade‐off between shade avoidance and prey capture. These results were unexpected. Inadequate replication and atypical precipitation patterns were ruled out as possible explanations. The population studied here (unlike that of a different, but morphologically similar, species growing in a fire‐maintained pine grass–sedge savanna) did not exhibit the ability to respond to variation in competition from neighboring plants.     

Effects of longleaf pine planting density and other factors on stand structure and associated wildlife habitat

The primary objective of many longleaf pine (Pinus palustris) restoration programs is to enhance or restore habitat for wildlife dependent on herbaceous plant communities. Because herbaceous cover is inversely related to canopy cover, restoration programs often place restrictions on longleaf pine planting density. However, the influence of planting density on understory plant communities has been inadequately evaluated. Therefore, we initiated a study to examine the relative influences of planting density and other factors on overall understory composition and forage availability for white‐tailed deer (Odocoileus virginianus) and northern bobwhite (Colinus virginianus) in nine longleaf pine stands throughout the Coastal Plain of Alabama during 2017–2018. We found that coverage of herbaceous plants decreased 3.5%, coverage of woody plants decreased 2.4%, and coverage of northern bobwhite forage plants decreased 1.9% for each 1 m²/ha increase in longleaf pine basal area. However, planting density was not a significant predictor of current basal area, nor coverage of any functional group of plants we examined, likely because current longleaf pine density averaged only 46% (range = 30–64%) of seedling planting density. We did not detect an effect of prescribed fire on stand condition or understory plant communities, likely due to variability in fire timing and frequency. Our findings related to planting density were likely a function of low longleaf pine survival, which is not uncommon. Because of this and the inherent variability in growth rates for young longleaf pine stands, restoration programs should consider placing greater emphasis on post‐planting monitoring and management than planting density.     

Declines in plant species richness and endemic plant species in longleaf pine savannas invaded by <Emphasis Type="Italic">Imperata cylindrica</Emphasis>

Despite the widespread perception that non-native species threaten biodiversity, there are few documented cases of non-native species displacing rare or specialized native species. Here, I examined changes in plant species composition over 5 years during patch expansion of a non-native grass, Imperata cylindrica, in longleaf pine flatwoods in Mississippi, USA. I used a multivariate approach to quantify the degree of habitat specialization and geographic range of all species encountered. I examined losses of species collectively as a function of plant height (controlling for initial frequency) and then the relationship between height and the degree of association with longleaf pine flatwoods, disturbed habitats, and the outer Gulf Coastal Plain of the southeastern USA. Patch expansion resulted in dramatic declines in species richness and increases in ground-level shade at both sites in just 3 years. Most tall saplings, shrubs, and vines were not endemic to longleaf pine communities and were less likely to be displaced than short herbs, most of which were indicative of longleaf pine communities. These results suggest that invasion of longleaf pine communities by I. cylindrica will likely cause significant losses of short, habitat-specialists and reduce the distinctiveness of the native flora of these threatened ecosystems.     

The Initial Phase of a Longleaf Pine‐Wiregrass Savanna Restoration: Species Establishment and Community Responses

The significant loss of the longleaf pine‐wiregrass ecosystem in the southeastern United States has serious implications for biodiversity and ecosystem functioning. In response to this loss, we have initiated a long‐term and landscape‐scale restoration experiment at the 80,125 ha (310 mi²) Department of Energy Savannah River Site (SRS) located near Aiken, South Carolina. Aristida beyrichiana (wiregrass), an important and dominant grass (i.e., a “matrix” species) of the longleaf pine savanna understory, and 31 other herbaceous “non‐matrix” species were planted at six locations throughout SRS in 2002 and 2003. Of the 36,056 transplanted seedlings, 75% were still alive in June 2004, while mean 1–2 year survival across all planted species was 48%. Lespedeza hirta (hairy lespedeza) exhibited the greatest overall survival per 3 × 3 m cell at 95%, whereas Schizachyrium spp. (little bluestem) exhibited the greatest mean cover among individual species at 5.9%. Wiregrass survival and cover were significantly reduced when planted with non‐matrix species. Aggregate cover of all planted species in restored cells averaged 25.9% in 2006. High rates of survival and growth of the planted species resulted in greater species richness (SR), diversity, and vegetative cover in restored cells. Results suggest that the loss of the longleaf pine‐wiregrass ecosystem may be ameliorated through restoration efforts and illustrate the positive impact of restoration plantings on biodiversity and vegetative cover.     

Historical Stem‐Mapped Permanent Plots Increase Precision of Reconstructed Reference Data in Ponderosa Pine Forests of Northern Arizona

Forest structural reference conditions are widely used to understand how ecosystems have been altered and guide restoration and management objectives. We used six stem‐mapped permanent plots established in the early twentieth century to provide precise structural reference conditions for ponderosa pine forests of northern Arizona prior to Euro‐American settlement. Reference conditions for these plots in 1873–1874 included the following historical attributes: tree densities of 45–127 trees/ha, mean tree diameter at breast height (dbh) of 43.8 cm with a corresponding quadratic mean diameter range of 41.5–51.3 cm, and a stand basal area of 9.2–18.0 m²/ha. The reconstructed diameter distributions (for live ponderosa pine trees with dbh ≥9.14 cm) prior to fire exclusion varied in shape but generally displayed an irregular unimodal distribution. We suggest that management objectives for the structural restoration of ponderosa pine forests of northern Arizona emphasize: (1) conservation and retention of all pre‐settlement (>130 years) trees; (2) reduction of tree densities with a restoration objective ranging between 50 and 150 trees/ha having a large‐tree component between 25 and 50% of the total trees per hectare, respectively; (3) manipulation of the diameter distribution to achieve a unimodal or irregular, uneven‐aged shape (possibly targeting a balanced, uneven‐aged shape on cinder soil types) through the use of harvest and thinning practices that mimic gap disturbances (i.e., individual tree selection system); and (4) retention of 3–11 snags and logs per hectare resulting from natural mortality.     

Habitat selection by northern spotted owls in mixed-coniferous forests

Conservation planning for the federally threatened northern spotted owl (Strix occidentalis caurina) requires an ability to predict their responses to existing and future habitat conditions. To inform such planning we modeled habitat selection by northern spotted owls based upon fine-scale (approx. 1.0 ha) characteristics within stands comprised primarily of mixed-aged, mixed coniferous forests of southwestern Oregon and north-central California. We sampled nocturnal (i.e., primarily foraging) habitat use by 71 radio-tagged spotted owls over 5 yr in 3 study areas and sampled vegetative and physical environmental conditions at inventory plots within 95% utilization distributions of each bird. We compared conditions at available forest patches, represented by the inventory plots, with those at patches used by owls using discrete-choice regressions, the coefficients from which were used to construct exponential resource selection functions (RSFs) for each study area and for all 3 areas combined. Cross-validation testing indicated that the combined RSF was reasonably robust to local variation in habitat availability. The relative probability that a fine-scale patch was selected decreased nonlinearly with distances from nests and streams; varied unimodally with increasing average diameter of coniferous trees and also with increasing basal area of Douglas-fir (Pseudotsuga menziesii) trees; increased linearly with increasing basal areas of sugar pine (Pinus lambertiana) and hardwood trees and with increasing density of understory shrubs. Large-diameter trees (>66 cm) appeared important <400 m from nest sites. The RSF can support comparative risk assessments of the short- versus long-term effects of silvicultural alternatives designed to integrate forest ecosystem restoration and habitat improvement for northern spotted owls. Results suggest fine-scale factors may influence population fitness among spotted owls. © 2011 The Wildlife Society.     

Predators and competitors of the mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae) in stands of changing forest composition associated with elevation

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Forest ecosystems of an Arizona Pinus ponderosa landscape: multifactor classification and implications for ecological restoration

Aim We developed an ecosystem classification within a 110,000‐ha Arizona Pinus ponderosa P. & C. Lawson (ponderosa pine) landscape to support ecological restoration of these forests. Specific objectives included identifying key environmental variables constraining ecosystem distribution and comparing plant species composition, richness and tree growth among ecosystems. Location The Coconino National Forest and the Northern Arizona University Centennial Forest, in northern Arizona, USA. Methods We sampled geomorphology, soils and vegetation on 66 0.05‐ha plots in open stands containing trees of pre‐settlement (c. 1875) origin, and on 26 plots in dense post‐settlement stands. Using cluster analysis and ordination of vegetation and environment matrices, we classified plots into ecosystem types internally similar in environmental and vegetational characteristics. Results We identified 10 ecosystem types, ranging from dry, black cinders/Phacelia ecosystems to moist aspen/Lathyrus ecosystems. Texture, organic carbon and other soil properties reflecting the effects of parent materials structured ecosystem distribution across the landscape, and geomorphology was locally important. Plant species composition was ecosystem‐specific, with C₃Festuca arizonica Vasey (Arizona fescue), for instance, abundant in mesic basalt/Festuca ecosystems. Mean P. ponderosa diameter increments ranged from 2.3–4.3 mm year^?1 across ecosystems in stands of pre‐settlement origin, and the ecosystem classification was robust in dense post‐settlement stands. Main conclusions Several lines of evidence suggest that although species composition may have been altered since settlement, the same basic ecosystems occurred on this landscape in pre‐settlement forests, providing reference information for ecological restoration. Red cinders/Bahia ecosystems were rare historically and > 30% of their area has been burned by crown fires since 1950, indicating that priority could be given to restoring this ecosystem's remaining mapping units. Ecosystem classifications may be useful as data layers in gap analyses to identify restoration and conservation priorities. Ecosystem turnover occurs at broad extents on this landscape, and restoration must accordingly operate across large areas to encompass ecosystem diversity. By incorporating factors driving ecosystem composition, this ecosystem classification represents a framework for estimating spatial variation in ecological properties, such as species diversity, relevant to ecological restoration.     

Alteration and Recovery of Slash Pile Burn Sites in the Restoration of a Fire‐Maintained Ecosystem

Restoration practices incorporating timber harvest (e.g. to remove undesirable species or reduce tree densities) may generate unmerchantable wood debris that is piled and burned for fuel reduction. Slash pile burns are common in longleaf pine ecosystem restoration that involves hardwood removal before reintroduction of frequent prescribed fire. In this context, long‐lasting effects of slash pile burns may complicate restoration outcomes due to unintended alterations to vegetation, soils, and the soil seed bank. In this study, our objectives were to (1) examine alterations to the soil seed bank, soil physical and chemical characteristics, and initial vegetation recolonization following burn and (2) determine the rate of return of soil and vegetation characteristics to pre‐burn conditions. We found that burning of slash piles (composed of scores of whole trees) results in elevated nutrient levels and significant impacts on vegetation and the soil seed bank, which remain evident for at least 6 years following burn. In this ecosystem, formerly weakly acidic soils become neutral to basic and levels of P remain significantly higher. Following an initial decrease after burn, total soil N increases with time since burn. These changes suggest that not only does pile burning create a fire scar initially devoid of biota, but it also produces an altered soil chemical environment, with possible consequences for long‐term ecosystem restoration efforts in landscapes including numerous fire scars. To facilitate restoration trajectories, further adaptive management to incorporate native plant propagules or suppress encroaching hardwoods within fire scars may be warranted in fire‐dependent ecosystems.