The effect of shelterwood harvesting and site preparation on eastern red-backed salamanders in white pine stands

We studied the effects of the regeneration cut of the shelterwood system and four site preparation options on populations of eastern red-backed salamanders in 90–100-year-old white pine forests in central Ontario, Canada. We established the study in 1994 using a randomized complete block design with three replicates and five treatments: (1) no harvest, no site preparation; (2) harvest, no site preparation; (3) harvest, mechanical site preparation; (4) harvest, chemical site preparation; (5) harvest, mechanical and chemical site preparation. We applied harvest and site preparation treatments from fall 1995 to fall 1997. We collected pre-treatment data in spring and summer of 1995 and post-treatment data from 1998 to 2002. We monitored salamander abundance using a grid of 20 cover boards surveyed 10 times per year within each of the 15 treatment plots. We also quantified changes in overstory and understory cover, supply of downed woody debris, and disturbance to the forest floor. Our data suggest that shelterwood cutting and site preparation can have immediate negative effects on the abundance of red-backed salamander populations in pine forest. However, effects are relatively short lived (<5 years). Changes in abundance appeared to be related to overstory and understory cover, and forest floor disturbance.     

Influence of coarse woody debris on herpetofaunal communities in upland pine stands of the southeastern Coastal Plain

Coarse woody debris (CWD) is thought to benefit herpetofauna in a variety of ways including serving as feeding sites, providing a moist environment, and providing protection from temperature extremes. We investigated the importance of CWD to amphibian and reptile communities in managed upland pine stands in the southeastern United States Coastal Plain during years 6 and 7 of a long-term study. Using a randomized complete block design, 1 of the following treatments was assigned to 9.3-ha plots: removal (n = 3; all downed CWD ≥10 cm in diameter and ≥60 cm long removed), downed addition (n = 3; five-fold increase in volume of down CWD), snag (n = 3; 10-fold increase in volume of standing dead CWD), and control (n = 3; unmanipulated). Herpetofauna were captured seasonally using drift-fence pitfall trapping arrays within treatment plots. We compared abundance, diversity, and richness of anurans, salamanders, lizards, and snakes using analysis of covariance with topographic variables (slope, elevation, aspect, and distance to nearest stream) included as covariates. We captured 355 amphibians and 668 reptiles seasonally from January 2007 to August 2008. Abundance, species richness, and species diversity were similar among treatments for anurans, salamanders, and lizards. Snake abundance, species richness, and diversity were higher in removal than downed addition plots. Anuran abundance increased as distance to nearest stream decreased. The majority of species captured during this study are adept at burrowing into the sandy soils of the region. Lack of reliance on CWD may be the result of herpetofaunal adaptation to the longleaf pine (Pinus palustris) ecosystem that historically dominated the upland areas of the study area.     

Woodland salamander response to two prescribed fires in the central Appalachians

Using coverboard arrays, we monitored woodland salamanders on the Fernow Experimental Forest in the central Appalachian Mountains, West Virginia, USA prior to and following two prescribed fires in mixed oak (Quercus spp.) forest stands. Treatments were burn plots on upper slopes or lower slopes fenced to prevent white-tailed deer (Odocoileus virginianus) herbivory or control plots that were unfenced and unburned. Most of the 7 species we observed were the mountain dusky salamander (Desmognathus ocropheaus), red-backed salamander (Plethodon cinereus) and slimy salamander (Plethodon glutinosis). Significant population responses were difficult to interpret with numerous treatment and year interactions. Results largely were equivocal. We found no change in woodland salamander assemblage prior to burning or afterwards. There were few differences in adult to juvenile ratios of salamanders among treatments. Still, a priori contrasts of mountain dusky salamanders and red-backed salamander counts corrected for detection probability were greater under coverboards in the 2 years monitored after both prescribed fires had occurred than before burning or in unburned controls. This suggests that these species responded to the reduced leaf litter on the forest floor by utilizing coverboards more. Similarly, the three predominate species of salamanders also were more numerous under coverboards in plots subjected to deer herbivory with less subsequent forest floor vegetation as compared to those burned plots that were fenced. Our observations would suggest that woodland salamanders somewhat are tolerant of two prescribed fires within close temporal proximity. However, because woodland salamanders can be significantly reduced following timber harvest, continued research is needed to fully understand impacts of fire as a pre-harvest management tool in central Appalachian forests.     

Amphibian and reptile community response to coarse woody debris manipulations in upland loblolly pine (Pinus taeda) forests

Coarse woody debris (CWD) has been identified as a key microhabitat component for groups that are moisture and temperature sensitive such as amphibians and reptiles. However, few experimental manipulations have quantitatively assessed amphibian and reptile response to varying CWD volumes within forested environments. We assessed amphibian and reptile response to large-scale, CWD manipulation within managed loblolly pine stands in the southeastern Coastal Plain of the United States from 1998 to 2005. Our study consisted of two treatment phases: Phase I treatments included downed CWD removal (removal of all downed CWD), all CWD removal (removal of all downed and standing CWD), pre-treatment snag, and control; Phase II treatments included downed CWD addition (downed CWD volume increased 5-fold), snag addition (standing CWD volume increased 10-fold), all CWD removal (all CWD removed), and control. Amphibian and anuran capture rates were greater in control than all CWD removal plots during study Phase I. In Phase II, reptile diversity and richness were greater in downed CWD addition and all CWD removal than snag addition treatments. Capture rate of Rana sphenocephala was greater in all CWD removal treatment than downed CWD addition treatment. The dominant amphibian and snake species captured are adapted to burrowing in sandy soil or taking refuge under leaf litter. Amphibian and reptile species endemic to upland southeastern Coastal Plain pine forests may not have evolved to rely on CWD because the humid climate and short fire return interval have resulted in historically low volumes of CWD.     

Structural changes and potential vertebrate responses following simulated partial harvesting of boreal mixedwood stands

Partial harvesting, where different numbers and arrangements of live trees are retained in forest stands, has been proposed for maintaining late-successional structure and associated vertebrate species within managed boreal forests. Using the stand dynamics model SORTIE-ND, we examined 80-year patterns of structural change in response to different intensities (30-70% basal area removal) and spatial patterns (22-273 m² mean patch size) of harvesting. We also applied habitat models for seven late-successional vertebrates to the structural conditions present after harvesting to assess potential species responses.Partial harvesting increased understory and downed woody debris (DWD) cover and decreased overstory structure for the first 25 years after harvest, in comparison to unharvested stands, with this effect subsequently reversing as harvest-induced regeneration reached the canopy. Although harvesting enhanced long-term structural development in this regard, large trees, large snags, and large DWD all remained below unharvested levels throughout the simulation period. Harvesting also produced transient increases in early-decay DWD and ground exposure. Most changes in structural attributes increased in proportion to harvest intensity, but structural differences among harvest patterns were generally small. Dispersed harvesting induced somewhat less pronounced decreases in vertical structure, and produced more post-harvest slash, than aggregated harvesting.All seven vertebrate species decreased in abundance as harvest intensity increased from 30 to 70%. In comparison to their pre-harvest abundances in old stands, vertebrates associated with DWD (redback salamander, marten, red-backed vole) showed neutral or positive responses at one or more harvest intensities, whereas those associated with large trees and snags (brown creeper, flying squirrel) consistently exhibited substantial adverse impacts.     

Long-term partial cutting impacts on Desmognathus salamander abundance in West Virginia headwater streams

To understand long-term impacts of partial cutting practices on stream-dwelling salamanders in the central Appalachians, we examined pooled abundance of Desmognathus fuscus and D. monticola salamanders (hereafter Desmognathus) in headwater streams located within long-term silvicultural research compartments on the Fernow Experimental Forest, Tucker County, West Virginia. We sampled Desmognathus salamanders in 12 headwater streams within silvicultural research compartments that have been subjected to partial cutting for approximately 50 years. We used an information-theoretic approach to test five a priori models explaining partial cutting effects at the compartment-level on Desmognathus abundance and eight a priori models explaining stream reach-scale habitat effects on Desmognathus abundance. Our modeling efforts resulted in the selection of two competing models explaining partial cutting effects on Desmognathus abundance at the compartment-level. The VOLUME model, which incorporated cumulative timber volume harvested within compartments, received the greatest support and indicated that Desmognathus abundance was impacted negatively by increased timber volume removal. The second model, LASTDISTURB, incorporating the single variable of time since last harvest activity, indicated that Desmognathus abundance increased with time since last harvest at the compartment-level. For stream reach-scale habitat variables, the EMBEDDED model incorporating the percent of embedded substrate within streams, received the strongest support for explaining Desmognathus abundance. Our results suggest that long-term partial cutting suppresses Desmognathus abundance, possibly by increasing stream sedimentation and thereby reducing available cover for juvenile and adult salamanders. However, these practices do not appear to have threatened long-term persistence of Desmognathus in central Appalachian headwater streams.     

Soricid response to coarse woody debris manipulations in Coastal Plain loblolly pine forests

We assessed shrew (soricids) response to coarse woody debris (CWD) manipulations in managed upland loblolly pine (Pinus taeda) stands in the upper Coastal Plain of South Carolina over multiple years and seasons. Using a completely randomized block design, we assigned one of the following treatments to 12, 9.3-ha plots: removal (n = 3; all CWD ≥ 10 cm in diameter and ≥60 cm long removed), downed (n = 3; 5-fold increase in volume of down CWD), snag (n = 3; 12-fold increase in standing dead CWD), and control (n = 3; unmanipulated). Therein, we sampled shrews during winter, spring, and summer seasons, 2003–2005, using drift-fence pitfall arrays. During 1680 drift-fence plot nights we captured 253 Blarina carolinensis, 154 Sorex longirostris, and 51 Cryptotis parva. Blarina carolinensis capture rate was greater in control than in snag treatments. Sorex longirostris capture rate was lower in removal than downed and control plots in 2005 whereas C. parva capture rate did not differ among treatments. Overall, the CWD input treatments failed to elicit the positive soricid response we had expected. Lack of a positive response by soricid populations to our downed treatments may be attributable to the early CWD decay stage within these plots or an indication that within fire-adapted pine-dominated systems of the Southeast, reliance on CWD is less than in other forest types.     

Decaying wood and tree regeneration in the Acadian Forest of Maine,USA

We examined the effect of management history on the availability of decayed downed wood and the use of downed wood as a regeneration substrate in mixed-species stands in the Acadian Forest of Maine. Regeneration of red spruce (Picea rubens Sarg.), eastern hemlock (Tsuga canadensis (L.) Carr.), balsam fir (Abies balsamea L. Mill), and red maple (Acer rubrum L.) was quantified. Treatments included variants of selection cutting, commercial clearcutting (unregulated harvesting), and no harvesting for >50 years (reference). Area of wood substrate (wood ≥ Decay Class III and ≥10 cm on at least one end) was less in the commercial clearcut than in the reference; other treatments were not differentiated. Spruce and hemlock seedlings were found at higher densities on wood than paired forest floor plots of equal area, regardless of treatment. Conversely, fir and maple were less abundant on wood than forest floor plots in reference and selection treatments, but more or equally abundant on wood than forest floor plots in the commercial clearcut. These findings suggest that silvicultural treatment affects both the availability of decayed downed wood and seedling-substrate relationships, and that forest management in the Acadian Region should consider availability of downed woody material.     

Abundance and attributes of wildlife trees and coarse woody debris at three silvicultural systems study areas in the Interior Cedar-Hemlock Zone,British Columbia

Unmanaged cedar (Thuja plicata)-hemlock (Tsuga heterophylla) forests of the northern Interior Wetbelt of British Columbia support standing and dead trees with a variety of structural features that provide habitat for wildlife. We describe the pre-harvest abundance and characteristics of wildlife trees (standing trees with special characteristics that provide habitat for wildlife) and coarse woody debris (CWD) at three silvicultural systems trials in cedar-dominated stands, and the short-term effects of forest harvesting on the abundance and attributes of CWD. The treatments were clearcut, group retention (70% volume removal), group selection (30% volume removal), and unlogged control. We measured standing trees in 75 0.125-ha plots and CWD along 225 24-m transects, using a functional classification system to characterize habitat attributes of trees and logs. CWD assessments were repeated on the same transects after the harvest. The relationship between tree size and occurrence of habitat features was strong for both standing trees and logs. Each of the four major tree species in the study area was associated with specific habitat features that occurred more often in that species than in any other. Large concealed spaces at the bases of trees, providing den sites and escape cover, were associated with hybrid white spruce (Picea engelmannii × glauca). We suggest that these trees had originated on nurse logs that subsequently rotted away; if that supposition is correct, there may be shortages of these structures in future stands that originate from plantations. Forest harvesting had little effect on the volume of CWD, but did affect the decay class distribution, reduce the proportion of pieces having structural habitat attributes, and reduce piece lengths; these effects were generally proportional to the level of harvest removal. Partial-cut silvicultural systems have the potential to mitigate anticipated deficits in large wildlife trees and logs in managed stands, if components of the stand are managed on longer rotations than those planned for timber production alone.     

Biofuel harvests, coarse woody debris, and biodiversity - A meta-analysis

Forest harvest operations often produce large amounts of harvest residue which typically becomes fine (foliage, small limbs and trees) and coarse woody debris (snags and downed logs). If removed at harvest, residual biomass has potential to be a local energy source and to produce marketable biofuel feedstock. But, CWD in particular serves critical life-history functions (e.g., breeding, foraging, basking) for a variety of organisms. Unfortunately, little is known about how forest biodiversity would respond to large scale removal of harvest residues. We calculated 745 biodiversity effect sizes from 26 studies involving manipulations of CWD (i.e., removed or added downed woody debris and/or snags). Diversity and abundance of both cavity- and open-nesting birds were substantially and consistently lower in treatments with lower amounts of downed CWD and/or standing snags, as was biomass of invertebrates. However, cumulative effect sizes for other taxa were not as large, were based on fewer studies, and varied among manipulation types. Little is currently known about biodiversity response to harvest of fine woody debris. Predicting the effects of biomass harvests on forest biodiversity is uncertain at best until more is known about how operational harvests actually change fine and coarse woody debris levels over long time periods. Pilot biomass harvests report post-harvest changes in CWD levels much smaller than the experimental changes involved in the studies we analyzed. Thus, operational biomass harvests may not change CWD levels enough to appreciably influence forest biodiversity, especially when following biomass harvest guidelines that require leaving a portion of harvest residues. Multi-scale studies can help reduce this uncertainty by investigating how biodiversity responses scale from the small scale of manipulative experiments (i.e., 10-ha plots) to operational forest management and how biodiversity response to CWD levels might vary at different spatial and temporal scales and in different landscape contexts.     

Tree size distribution and abundance explain structural complexity differentially within stands of even-aged and uneven-aged structure types

Characterizations of physical structural complexity are an important surrogate for the potential of forested stands to provide desired ecosystem services such as biodiversity. Distinguishing between stands with different structural conditions is not only a necessary feature of useful structural metrics and indices, but how such measures vary among stands can reveal clues to the ecological processes driving structure. We used stand inventory metrics and indices of structural complexity to differentiate between even-aged and uneven-aged structure types using 10 stem-mapped coniferous stands of each type distributed across Switzerland. Within each structure type, we further explored relationships among stand inventory metrics and structural indices over a roughly 10-year period of management intervention. The even-aged and uneven-aged structure types were clearly differentiated using both stand inventory metrics and spatially explicit structural complexity indices. Overall, structural complexity within even-aged stands was strongly related to, and best predicted by, metrics including the distribution of basal area among canopy layers, while complexity in the uneven-aged stands was most strongly related to, and best predicted by, metrics including measures of abundance. Although predictive models could be developed for canopy position mixture, diameter differentiation, and small-scale structural complexity (but not spatial aggregation) using only stand inventory metrics, the prediction success after only a single management intervention was lower than expected. These results indicate that research to explore small-scale structural complexity requires detailed spatially explicit inventory data and that management to enhance structural complexity may require the manipulation of different attributes in stands of even-aged (diameter distribution) and uneven-aged (total abundance) structure types.     

Efficacy of riparian buffers in mitigating local population declines and the effects of even-aged timber harvest on larval salamanders

Headwater streams are an important and prevalent feature of the eastern North American landscape. These streams provide a wealth of ecosystem services and support tremendous biological diversity, which is predominated by salamanders in the Appalachian region. Salamanders are ubiquitous throughout the region, contributing a significant biomass that supports ecological and ecosystem processes. One of the greatest threats to salamanders is loss of headwater-riparian habitat through timber harvest. In this study, we measured larval salamander abundance at five headwater streams with different riparian buffer widths retained following logging. By sampling larval salamanders using leaf litter bags, we assessed the impacts of even-aged timber harvest on aquatic larval salamander abundances, where it was found that larvae are negatively impacted by increased stream sedimentation and a decrease in riparian buffer width. We found that retention of a 9-m buffer was effectively no different than complete removal of all riparian forest, and as such, current regulations to protect headwater streams are ineffectual. Furthermore, no significant differences were observed between the 30 m buffer treatment and uncut control treatments suggesting that a 30 m or larger riparian buffer may assuage the in-stream effects of riparian timber harvest. Management guidelines for Appalachian forests should be revised to accommodate the biology of plethodontid salamanders.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Dead wood in clearcuts of semi-natural forests in Estonia: site-type variation, degradation, and the influences of tree retention and slash harvest

Semi-natural forests, which naturally regenerate after timber harvesting, provide distinct opportunities for dead wood (DW) management for biodiversity. We described DW pool and sources of its variation during the first decade after final felling in Estonia, hemiboreal Europe. Depending on forest type, the mean post-harvest volumes of above-ground DW ranged from 70 to 119 m³ ha^?1. Final felling generally did not reduce downed coarse woody debris (CWD) because many sawn logs were left on-site, and soil scarification was rarely used. However, subsequent decay of downed CWD appears to be accelerated due to the increased ground contact of logs, so that even the relatively small inputs from live retention trees observed (5 m³ ha^?1 per decade) can be ecologically significant. While final felling greatly reduced snag abundance, the mortality of retained live trees generally balanced their later losses. The volumes of downed fine woody debris in conventional cutover sites were roughly double that of pre-harvest forests. Slash harvest caused an approximately twofold reduction in downed DW and resulted in CWD volumes that were below mature-forest levels. The results indicate that the habitat quality of cutovers critically depends both on the retention and on the post-harvest management of biological legacies. In Estonia, the necessary improvements include more careful retention of snags in final felling, selecting larger retention trees, focusing slash harvest on the fine debris of common tree species, and providing snags of late-successional tree species.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

Influence of fallen coarse woody debris on the diversity and community structure of forest-floor spiders (Arachnida: Araneae)

Purely observational studies have documented differences in the abundance and diversity of several litter-dwelling arthropods between sites adjacent to, and far from, CWD, which suggests that reduction of fallen coarse woody debris (CWD) in temperate forests by traditional forestry practices may affect the litter arthropod community. As few field experiments have directly tested the impact of CWD on arthropods inhabiting the litter at different distances from CWD, we removed CWD from replicated open plots on the floor of a second-growth deciduous forest in order to reveal the causal connection between CWD and litter-dwelling spiders, often the most diverse and abundant predators among the litter macrofauna of temperate forests. We also documented the impact of the CWD manipulation on spider prey and several other major macroarthropod groups. Before removing CWD (∅ = 14.3 ± 0.7 cm), we measured response variables as a function of distance (0.5–1.5 m) from CWD in both removal and control plots. In agreement with results of previous research that solely utilized this observational approach in temperate forests, volume and dry mass of litter, spider diversity, overall spider density, and densities of 8 of 16 major spider genera were higher adjacent to CWD before experimental manipulations. Removing CWD reduced the amount of litter and the density of spiders in litter close to where the CWD had been. Removing CWD also altered spider community structure, which had differed between litter sites adjacent to, and far from, CWD prior to the experimental removal of CWD. The patterns, though, were not completely congruent, as some of the taxa affected by the manipulation had not differed between sites prior to the removal of CWD, and vice-versa. Our findings suggest that complex interactions among structural, biotic and microclimatic factors underlie the observed responses to CWD removal by spiders and other arthropods in the litter layer. We also conclude that drawing inferences solely from observational studies is not a reliable approach for predicting the impact of changes in the amount of CWD on arthropods of forest-floor leaf litter. Further field experiments manipulating different volumes of CWD are needed in order to determine the minimum amount of CWD that should be kept on the forest floor of managed forests in order to maintain densities and diversities of major leaf-litter arthropods.     

Three-dimensional light structure of an upland Quercus stand post-tornado disturbance

Light is the most common limiting factor in forest plant communities,influencing species composition,stand structure,and stand productivity in closed canopy stands.Stand vertical light structure is relatively simple under a closed canopy because most light is captured by overstory trees.However,wind disturbance events create canopy openings from local to landscape scales that increase understory light intensity and vertical light structural complexity.We studied the effects of an EF-1 tornado on horizontal and vertical(i.e.three-dimensional)light structure within a Quercus stand to determine how light structure changed with increasing disturbance severity.We used a two-tiered method to collect photosynthetic photon flux density at 4.67 m and 1.37 m above the forest floor to construct three-dimensional light structure across a canopy disturbance severity gradient to see if light intensity varied with increasing tornado damage.Results indicate that increased canopy disturbance closer to the tornado track increased light penetration and light structure heterogeneity at lower forest strata.Increased light intensity correlated with increased sapling density that was more randomly distributed across the plot and had shifted light capture higher in the stand structure.Light penetration through the overstory was most strongly correlated with decreased stem density in the two most important tree species(based on relative dominance and relative density)in the stand,Quercus alba L.(r=-0.31)and Ostrya virginiana(Mill.)K.Koch(r=-0.27,p<.01),and indicated that understory light penetration was most affected by these two species.As managers are increasingly interested in patterning silvicultural entries on natural disturbances,they must understand residual stand and light structures that occur after natural disturbance events.By providing spatial light data that quantifies light structure post-disturbance,managers can use these results to improve planning required for long-term management.The study also provides comparisons with anthropogenic disturbances to the midstory that may offer useful comparisons to natural analogs for future silvicultural consideration.     

Effects of even-aged timber harvest on stream salamanders: Support for the evacuation hypothesis

Habitats worldwide are increasingly threatened by degradation and conversion. Critical to the process of habitat loss is the organismal response, which can have effects on immediate conservation measures or future restoration. Among the most threatened and underappreciated habitats are headwater streams, which are small but abundant features of montane forests. These habitats comprise a significant proportion of the total stream length, can harbor remarkable biodiversity, and are critical for numerous ecosystem processes. One of the most abundant organisms in montane headwater ecosystems are salamanders, and therefore what happens to salamanders when the forest habitats surrounding headwater streams are altered? Three main hypotheses exist: (1) mortality hypothesis; (2) retreat hypothesis; and (3) evacuation hypothesis. To examine these hypotheses we evaluated the impacts of even-aged riparian timber harvest on stream-breeding salamanders. Riparian forests along headwater streams were logged, leaving riparian buffers of 0 m, 9 m, and 30 m. Responses to each riparian alteration were measured in terms of salamander terrestrial habitat use and growth in the riparian habitat, as well as changes in population density within headwater streams. Adult and juvenile salamander densities measured in headwater streams were significantly greater in logged riparian treatments than in unaltered riparian treatments. In addition, salamanders significantly reduced their terrestrial habitat use following riparian logging with both the average distance from the stream and the relative abundance of salamanders decreasing. It is unlikely that salamanders will persist in highly modified riparian habitats, as we measured significantly reduced body conditions over short periods of time at these sites. We present corroborative evidence that salamanders evacuate the riparian habitat following intensive riparian logging, emigrating to adjacent headwater streams. Our results underscore the sensitivity of stream salamanders to riparian habitat alteration as well as the importance of riparian buffers in preserving amphibian assemblages.     

Modeling production and decay of coarse woody debris in loblolly pine plantations

Forest management can have large impacts on the production and yield of coarse woody debris (CWD) in terrestrial ecosystems, yet few modeling tools exist to inform such efforts. The goal here was to develop a set of prediction equations for use in conjunction with loblolly pine (Pinus taeda L.) modeling and inventory systems to estimate CWD yields at scales ranging from individual trees to whole plantations. Permanent field plots from a 21-year study of thinning effects on plantation growth and yield across the commercial range of the species in the southern United States were surveyed to obtain sample data on CWD volume, density, and mass. Measured CWD properties were combined with inventory records of tree mortality over the study duration to characterize CWD production, decay and yield in a series of prediction equations. The resulting equations predict CWD attributes of dead trees including dry weight (kg) and fraction of standing versus downed woody material based on the time since death (years), tree diameter at breast height (cm) and height (m) at time of death and geographic coordinates of latitude and longitude. A stand-level equation predicts total CWD yield (Mg ha⁻¹) for thinned or unthinned stands based on plantation age, stem density (trees ha⁻¹), and the average height of dominant and codominant trees (m). Piece-level equations predict dry density (kg m⁻³) or nitrogen concentration (%) of CWD pieces based on their position (standing or down), ordinal decay classes, and latitude. The tree and stand-level prediction equations are designed for use in GIS or growth and yield modeling systems. The piece-level equations are designed to be used in inventory applications that survey CWD. The equations should facilitate the accurate and facile determination of mass, carbon, and nitrogen contents of CWD in planted loblolly pine forests of the southern United States.