Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Bark beetle community structure under four ponderosa pine forest stand conditions in northern Arizona

We studied the bark beetle guild (Coleoptera: Scolytidae) in the ponderosa pine forests of northern Arizona to explore if the species assemblages and relative abundance differ between managed and unmanaged stands. Four stand conditions were assessed: (1) unmanaged stands with high tree density, (2) thinned stands, (3) thinned and burned (with prescribed fire) stands and (4) stands that had been burned by stand replacing wildfires. The study was conducted in the ponderosa pine forests of the Coconino Plateau, northern Arizona. For several decades this area has been relatively free of bark beetle outbreaks despite the current overstocked condition of many stands. We found that a similar species assemblage composed of Dendroctonus frontalis, D. brevicomis, D. valens, D. approximatus, D. ponderosae, and Ips pini occurred across all four stand conditions over 3 years of study. The population levels of all these species were endemic across all stand conditions. The non-aggressive D. approximatus and D. valens were indicator species for thinned and unmanaged stands, respectively, but this was not consistent among years. The ambrosia beetle Gnathotrichus sp. and the bark beetle predator Enoclerus sp. consistently indicated stands burned by wildfire. In addition to our field experiment, we analyzed the historical pattern of attacks of bark beetles in our area of study. Our findings suggest that the pattern of attack of D. brevicomis (the only Dendroctonus species for which attacks have been reported) and Ips spp. has been through scattered small infestations in groups of 1–10 trees. Whereas small infestations by Ips spp. are increasing, those for D. brevicomis are decreasing. Although we agree that the high density stands in northern Arizona are in an “unhealthy” condition, our results do not show that they were supporting large bark beetle outbreaks. Our results challenge the theoretical assumptions about the relationship between stand structure, tree resistance and bark beetle performance.     

Perpetuating old ponderosa pine

We review current knowledge about the use of management treatments to reduce human-induced threats to old ponderosa pine (Pinus ponderosa) trees. We address the following questions: Are fire-induced damage and mortality greater in old than younger trees? Can management treatments ameliorate the detrimental effects of fire, competition-induced stress, and drought on old trees? Can management increase resistance of old trees to bark beetles? We offer the following recommendations for the use of thinning and burning treatments in old-growth ponderosa pine forests. Treatments should be focused on high-value stands where fire exclusion has increased fuels and competition and where detrimental effects of disturbance during harvesting can be minimized. Fuels should be reduced in the vicinity of old trees prior to prescribed burns to reduce fire intensity, as old trees are often more prone to dying after burning than younger trees. Raking the forest floor beneath old trees prior to burning may not only reduce damage from smoldering combustion under certain conditions but also increase fine-root mortality. Thinning of neighboring trees often increases water and carbon uptake of old trees within 1 year of treatment, and increases radial growth within several years to two decades after treatment. However, stimulation of growth of old trees by thinning can be negated by severe drought. Evidence from young trees suggests that management treatments that cause large increases in carbon allocation to radial xylem growth also increase carbon allocation to constitutive resin defenses against bark beetle attacks, but evidence for old trees is scarce. Prescribed, low-intensity burning may attract bark beetles and increase mortality of old trees from beetle attacks despite a stimulation of bole resin production.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

Effects of prescribed fire and season of burn on direct and indirect levels of tree mortality in Ponderosa and Jeffrey Pine Forests in California,USA

Many forests that historically experienced frequent low-intensity wildfires have undergone extensive alterations during the past century. Prescribed fire is now commonly used to restore these fire-adapted forest ecosystems. In this study, we examined the influence of prescribed burn season on levels of tree mortality attributed to prescribed fire effects (direct mortality) and bark beetles (Coleoptera: Curculionidae, Scolytinae) (indirect mortality) in ponderosa pine, Pinusponderosa Dougl. ex Laws., and Jeffrey pine, Pinusjeffreyi Grev. and Balf., forests in California, USA. A total of 816 trees (9.9% of all trees) died during this 3-yr study. Significantly higher levels of tree mortality (all sources) occurred following early and late season burns compared to the untreated control, but no significant difference was observed between burn treatments. The majority (461 trees) of tree deaths were attributed to direct mortality from prescribed burns and was strongly concentrated (391 trees) in the smallest diameter class (<20.2 cm diameter at breast height, dbh). For the largest trees (>50.7 cm dbh), significantly higher levels of tree mortality occurred on early season burns than the untreated control, most of which resulted from indirect mortality attributed to bark beetle attacks, specifically western pine beetle, Dendroctonus brevicomis LeConte, and mountain pine beetle, D. ponderosae Hopkins. Red turpentine beetle, D. valens LeConte, was the most common bark beetle species found colonizing trees, but tree mortality was not attributed to this species. A total of 355 trees (4.3% of all trees) were killed by bark beetles. Dendroctonus brevicomis (67 trees, 18.9%) and D. ponderosae (56 trees, 15.8%), were found colonizing P. ponderosa; and Jeffrey pine beetle, D. jeffreyi Hopkins, was found colonizing P. jeffreyi (seven trees, 2.0%). We also found pine engraver, Ips pini (Say) (137 trees, 38.6%), and, to a much lesser extent, Orthotomicus (=Ips) latidens (LeConte) (85 trees, 23.9%) and emarginate ips, I. emarginatus (LeConte) (3 trees, 0.8%) colonizing P. ponderosa and P. jeffreyi. Few meaningful differences in levels of indirect tree mortality attributed to bark beetle attack were observed between early and late season burns. The incidence of root and root collar pathogens (Leptographium and Sporothrix spp.), including species known to be vectored by bark beetles, was low (18% of trees sampled). The implications of these and other results to management of P. ponderosa and P. jeffreyi forests are discussed in detail.     

Tree mortality in drought-stressed mixed-conifer and ponderosa pine forests, Arizona, USA

We monitored tree mortality in northern Arizona (USA) mixed-conifer and ponderosa pine (Pinus ponderosa Dougl. ex Laws) forests from 1997 to 2007, a period of severe drought in this area. Mortality was pervasive, occurring on 100 and 98% of 53 mixed-conifer and 60 ponderosa pine plots (1-ha each), respectively. Most mortality was attributable to a suite of forest insects, mediated by drought stress. The number of trees dying from 2002 to 2007 was more than 200% greater than the number dying from 1997 to 2002 in mixed-conifer forest and 74% greater in ponderosa pine forest. Extent of mortality was spatially variable in both forest types. Median cumulative mortality (the ratio of dead to live trees) increased by approximately 53 and 65% in mixed-conifer and ponderosa pine forests, respectively, from 2002 to 2007. Median mortality rates from 2002 to 2007 were approximately 2.0% year⁻¹ in mixed-conifer forest (range = 0-28.5%) and 0.4% year⁻¹ in ponderosa pine forest (range = 0-13.6%). Mortality rates generally were not strongly related to either elevation or stand density. Mortality was nonrandom with respect to tree size classes and species. Proportions of trees dying were greatest in the largest size classes, particularly in mixed-conifer forest, where mortality in the largest size class exceeded 22% from 2002 to 2007. Mortality in mixed-conifer forest was particularly pronounced for quaking aspen (85%) and white fir (28%), the least drought tolerant species present. These results provide an early glimpse of how these forest types are likely to respond to predicted climate changes in the southwestern USA. They suggest that these forests are not resilient to climate change, and that treatments to increase resilience to climate change may be appropriate. Research on causes of spatial heterogeneity in extent of mortality might suggest valuable approaches to aid in increasing resilience.     

Recruitment of ponderosa pine seedlings in the Cascade Range

Abiotic stresses on seedling regeneration in xeric ecosystems are great, hence recruitment processes can be facilitated by stand factors that ameliorate the germinant-scale microenvironment. An experiment was conducted on the eastern slope of the Cascade Range to test the effects of shrub cover, simulated seed caching, and substrate on the recruitment of Pacific ponderosa pine (Pinus ponderosa var. ponderosa Dougl. ex Laws.) seedlings. Failure rates of seeds sowed in exclosures were large, with less than 30% emerging as germinants in the spring following fall sowing. Simulated seed caching improved emergence rates by more than sevenfold and was responsible for 88% of all spring germinants. Emergence rates were lowest from uncached seeds on litter. Just 16% of the crop survived the summer and fall to the month of November, or less than 5 months after emergence. Shrub cover did not affect emergence rates, but establishment rates were higher: seedlings beneath shrubs succumbed to desiccation at a slower rate than unshaded seedlings. By August there were 2.3 times more survivors at shrub-shaded sites than unshaded sites, and by the end of fall, when seedlings were considered established, more than 78% existed beneath shrubs. This study provides evidence that the natural recruitment of ponderosa pine seedlings is facilitated by the occurrence of the species’ common shrub associates.     

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.     

Bark beetle (Coleoptera: Scolytidae) diversity in spaced and unmanaged mature lodgepole pine (Pinaceae) in southeastern British Columbia

Variation in the number and diversity of bark beetles in spaced mature lodgepole pine stands in the East Kootenay region of British Columbia was analyzed in relation to location (site), spacing treatment and years following treatment. We analyzed the number of bark beetles and the number of bark beetle species that emerged from stumps or were captured in flight traps in the first five years following spacing. We also investigated the incidence of bark beetle attacks on the remaining trees and the mean dates of emergence from stumps and of capture in flight traps for the common species. Observations were made on three sites, each having three treatments: 4 m × 4 m spacing, 5 m × 5 m spacing, and an untreated control. The mean density of bark beetles emerged from stumps was different among sites and years but not between spacing treatments. There was no statistically significant variation in the number of bark beetle species captured in flight traps by site, spacing treatment, years, or spacing treatment and years. Significantly more bark beetles were captured in the 4 m × 4 m spacing treatment than in the control. The number of bark beetles captured was the highest in the first 2 years following treatment. Up to 26 species of bark beetles, excluding ambrosia beetles, were captured in flight barrier traps. There was no difference in species diversity by site or treatment indicating that species diversity in mature lodgepole pine is relatively stable over large areas. Of the 213 trees that sustained at least 10 attacks by bark beetles on the lower 2 m of the bole, 59.1% occurred in the spaced plots but only 18.2% of those were successful, versus 74.7% success in the infested trees in the control plots. The majority of infested trees contained Ips sp., Dendroctonus valens and D. murrayanae. Of the seven trees attacked by mountain pine beetle (D. ponderosae) only one tree was located in a spaced plot.     

Risk rating for mountain pine beetle infestation of lodgepole pine forests over large areas with ordinal regression modelling

The mountain pine beetle Dendroctonus ponderosae Hopkins is endemic to lodgepole pine, Pinus contorta var. latifolia Engelmann, forests in western Canada. However, the current beetle epidemic in this area highlights the challenges faced by forest managers tasked with prioritizing stands for mitigation activities such as salvage harvesting and direct control methods. In western Canada, the operational risk rating system for mountain pine beetle is based on biological knowledge gained from a rich legacy of stand-scale field studies. Due to the large spatial (millions of hectares affected) and temporal (over 10 years) extents of the current epidemic, new research into large-area mountain pine beetle processes has revealed further insights into the landscape-scale characteristics of beetle infested forests. In this paper, we evaluated the potential for this new knowledge to augment an established system for rating the short-term risk of tree mortality in a stand due to mountain pine beetle. New variables explored for utility in risk rating include direct shortwave radiation, site index, diameter at breast height, the temporal trends in local beetle populations, Biogeoclimatic Ecosystem Classification and beetle–host interaction variables. Proportional odds ordinal regression was used to develop a model for the Vanderhoof Forest District in west-central British Columbia. Prediction on independent data was assessed with the area under the receiver operator curve (AUC), indicating good discriminatory power (AUC = 0.84) for predicting levels of mountain pine beetle-caused pine mortality.     

The effects of a thinning treatment on carbon stocks in a northern Arizona ponderosa pine forest

Vast areas of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forest in the western United States have become unnaturally dense because of relatively recent land management practices that include fire suppression and livestock grazing. In many areas, thinning treatments can re-establish the natural ecological processes and help restore ecosystem structure and function. Precipitous global climate change has focused attention on the carbon storage in forests. An unintended consequence of fire suppression has been the increased storage of carbon in ponderosa stands. Thinning treatments reduce standing carbon stocks while releasing carbon through the combustion of fuel in logging machinery, burning slash, and the decay of logging slash and wood products. These reductions and releases of stored carbon must be compared to the risk of catastrophic fire burning through the stand and releasing large quantities of carbon to the atmosphere to more fully understand the costs and benefits – in carbon terms – of forest restoration strategies.     

Post-1935 changes in forest vegetation of Grand Canyon National Park, Arizona, USA: Part 1 - ponderosa pine forest

Vegetation plots originally sampled in Grand Canyon National Park (GCNP), Arizona, USA in 1935 are the earliest-known, sample-intensive, quantitative documentation of forest vegetation over a Southwest USA landscape. These historical plots were located as accurately as possible and resampled in 2004 to document multi-decadal changes in never-harvested Southwestern forests. Findings for ponderosa pine forest (PPF) differed among three forest subtypes (dry, mesic, and moist PPF), indicating that understanding the ecology of PPF subtypes is essential for development of ecologically based management practices. Dry PPF, which is transitional with pinyon-juniper vegetation at low elevation, exhibited no changes from 1935 to 2004. Mesic PPF, the core subtype of PPF, had increased densities of total trees, ponderosa pine (Pinus ponderosa), and white fir (Abies concolor) in the 10-29.9 cm diameter class from 1935 to 2004 that may have induced decreased densities of larger ponderosa pines and total tree and ponderosa pine basal areas. Moist PPF, which is transitional with mixed conifer forest at high elevation, was the most dynamic PPF subtype with decreases from 1935 to 2004 in total density and total basal area that are largely attributable to decreases in quaking aspen (Populus tremuloides). Graphical synthesis of datasets with historical and modern values for density and basal area indicates that overall PPF (all subtypes combined) increased in sapling density of all species combined and conifers with canopy potential and decreased in density of quaking aspen trees since the late 19th century. PPF of GCNP has passed through an accretion phase of forest development with increases in density and, depending on PPF subtype and variable being examined, is at or past the point of inflection to recession of density and basal area. Increases in small diameter ponderosa pine and white fir from 1935 to 2004 portend potential additional accretion, but decreases in total basal area, density and basal area of quaking aspen, basal area of ponderosa pine, and density of larger diameter ponderosa pine indicate PPF has passed the inflection point from accretion to recession. Uncertainties about 19th-century PPF structure and composition and about future ecological and societal environments lead to the conclusion that resource managers of GCNP and other natural areas should consider a change in focus from the objective of achieving desired future conditions to an objective of avoiding undesired future conditions.     

Soil organic matter in a ponderosa pine forest with varying seasons and intervals of prescribed burn

Prescribed burning is used to reduce fuel loads and return ponderosa pine forests of the Western U.S. to their historical structure and function. The impact of prescribed burning on soil is dependent on fire severity which is largely managed by burning in the fall or the spring; frequency of fire will also regulate long-term fire impacts. The objective of this study was to determine if soils and soil organic matter (SOM) were affected by prescribed burning in the fall or the spring using singular or multiple prescribed burns. Prescribed burning was initiated in the spring of 1997 and fall of 1997 at 5-year intervals and once during a 15-year period on a study site located within the Malheur National Forest of the southern Blue Mountains of eastern Oregon. Soils were sampled by major genetic horizon in 2004. The 5-year interval plots had burned twice with 1–2 years of recovery while the 15-year interval plots had burned only once with 6–7 years of recovery. Samples were analyzed for pH, carbon (C), nitrogen (N), C/N ratio, cation exchange capacity, base saturation, water repellency, and humic substance composition by alkali extraction. Fall burning decreased C and N capital of the soil (O horizon +30 cm depth mineral soil) by 22–25%. Prescribed burning did not have an effect on fulvic or humic acid C concentration (FA and HA, respectively) of the mineral soil and only a minor effect on FA and HA concentration of the O horizon. One or two fall burns decreased humin and the alkali non-soluble C (NS) content of O horizon by 15 and 30%, respectively. Initiating fall burning in fire-suppressed stands may not preserve soil C, N, humin, and NS content, but may replicate the natural fire regime. Spring burning using a return interval of 5 or more years reduces the fuel load while having little impact on soil C, N, and SOM composition and may be used to prepare a site for subsequent fall burns.     

Effects of vertebrate herbivores and shrub characteristics on arthropod assemblages in a northern Arizona forest ecosystem

Large herbivores have potential to affect invertebrate community structure through numerous processes, but little work has been done to evaluate the relative importance of direct and indirect factors. In this study, we measured arthropod community assemblages on Ceanothus fendleri A. Gray (Fendler's ceanothus) plants that were growing inside and outside of 4-m² exclosures. We used univariate analyses and structural equation modeling (SEM) to evaluate relationships within this herbivore–plant–arthropod system in ponderosa pine (Pinus ponderosa Laws.) forests of northern Arizona, USA. Results showed that individual arthropod abundance, family diversity, family richness, and functional group richness were significantly greater on plots where C. fendleri plants were protected from large ungulate herbivores (e.g., mule deer (Odocoileus hemionus) and Rocky Mountain elk (Cervus elaphus nelsoni)) than on unprotected plots in each of the three study years. Results also indicated the following: (1) arthropod abundance was significantly greater on protected plants than unprotected plants; (2) rarefaction curves suggested arthropod family richness was similar between protected and unprotected plants in two of the three years when scaled by number of individuals but the estimated total richness was consistently higher on protected plants; (3) arthropod abundance was directly affected by protection from herbivores, plant stem length, and number of flowers; (4) arthropod family richness was related to the number of individuals collected and affected by stem length. Results from this study illustrate that arthropod communities are directly affected by foraging vertebrate herbivores as well as indirectly affected through complex plant-mediated factors in this model system. Protection of preferred forage plants such as C. fendleri from ungulate herbivores can potentially increase diversity of arthropod assemblages in these forests, help conserve biological diversity, and enhance ecosystem restoration efforts.     

Thinning reduces soil carbon dioxide but not methane flux from southwestern USA ponderosa pine forests

Forest soils are important components of the global carbon cycle because they both store and release carbon. Carbon dioxide is released from soil to the atmosphere as a result of plant root and microbial respiration. Additionally, soils in dry forests are often sinks of methane from the atmosphere. Both carbon dioxide and methane are greenhouse gases whose increasing concentration in the atmosphere contributes to climate warming. Thinning treatments are being implemented in ponderosa pine forests across the southwestern United States to restore historic forest structure and reduce the risk of severe wildfire. This study addresses how thinning alters fluxes of carbon dioxide and methane in ponderosa pine forest soils within one year of management and examines mechanisms of change. Carbon dioxide and methane fluxes, soil temperature, soil water content, forest floor mass, root mass, understory plant biomass, and soil microbial biomass carbon were measured before and after the implementation of a thinning and in an unthinned forest. Carbon dioxide efflux from soil decreased as a result of thinning in two of three summer months. Average summer carbon dioxide efflux declined by an average of 34 mg C m⁻² hr⁻¹ in the first year after thinning. Methane oxidation did not change in response to thinning. Thinning had no significant short-term effect on total forest floor mass, total root biomass, or microbial biomass carbon in the mineral soil. Understory plant biomass increased after thinning. Thinning increased carbon available for decomposition by killing tree roots, but our results suggest that thinning reduced carbon dioxide emissions from the soil because the reduction in belowground autotrophic respiration was larger than the stimulation of heterotrophic respiration. Methane oxidation was probably not affected by thinning because thinning did not alter the forest floor mass enough to affect methane diffusion from the atmosphere into the soil.     

Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

Wildlife responses to thinning and burning treatments in southwestern conifer forests: A meta-analysis

After a century of fire suppression, conifer forests in the western United States have dramatically departed from conditions that existed prior to Euro-American settlement, with heavy fuel loads and an increased incidence of wildfire. To reduce this threat and improve overall forest health, land managers are designing landscape-scale treatments that strategically locate thinning and burning treatments to disrupt fuel continuity, allowing managed wildfires to burn the remaining area. A necessary step in designing and evaluating these treatments is understanding their ecological effects on wildlife. We used meta-analysis to compare effects of small-diameter removal (thinnings and shelterwoods) and burning treatments, selective harvesting, overstory removal (including clearcutting), and wildfire on wildlife species in southwestern conifer forests. We hypothesized that small-diameter removal and burning treatments would have minimal effects on wildlife compared to other treatments. We found 33 studies that met our criteria by (1) comparing density or reproductive output for wildlife species, (2) using forest management or wildfire treatments, (3) implementing control-impact or before-after control-impact design using unmanaged stands as controls, and (4) occurring in Arizona or New Mexico ponderosa pine (Pinus ponderosa) or mixed conifer (Abies/Picea/Pinus) forest. The 22 studies suitable for meta-analysis occurred ≤20 years post-treatment on sites <400 ha. Small-diameter harvest and burning treatments had positive effects but thin/burn and selective harvest treatments had no detectable effect on most small mammals and passerine bird species reported in studies; overstory removal and wildfire resulted in an overall negative response. We examined foraging guild responses to treatments; ground-foraging birds and rodents had no strong response. Aerial-, tree-, and bole-foraging birds had positive or neutral responses to the small-diameter removal and burning treatments, but negative responses to overstory removal and wildfire. Small-diameter removal and burning treatments as currently being implemented in the Southwest do not negatively impact most of the wildlife species in the studies we examined in the short-term (≤10 years). We believe a combination of treatments in a patchy arrangement across the landscape will result in the highest diversity and density. We recommend that managers implement thinning and burning treatments, but that future research efforts focus on long-term responses of species at larger spatial scales, use reproductive output as a more informative response variable, and target species for which there is a paucity of data.     

Southern pine beetle regional outbreaks modeled on landscape, climate and infestation history

The southern pine beetle (Dendroctonus frontalis, SPB) is the major insect pest of pine species in the southeastern United States. It attains outbreak population levels sufficient to mass attack host pines across the landscape at scales ranging from a single forest stand to interstate epidemics. This county level analysis selected and examined the best climatic and landscape variables for predicting infestations at regional scales. The analysis showed that, for a given county, the most important factor in predicting outbreaks was that the county was classified as in outbreak status in the previous year. Other important factors included minimum winter temperature and the greatest difference between the average of daily minimums and a subsequent low temperature point, precipitation history either seasonally in the previous year or difference from average over the previous 2 years, the synchronizing effect of seasonal temperatures on beetle populations and the relative percentage of total forest area composed of host species. The statistical models showed that climatic variables are stronger indicators of outbreak likelihood than landscape structure and cover variables. Average climatic conditions were more likely to lead to outbreaks than extreme conditions, supporting the notion of coupling between a native insect and its native host. Still, some extreme events (i.e., periods of very low temperature or very high precipitation) did precede beetle infestation. This analysis suggested that there are predisposing and inciting factors at the large scale but the driving factors leading to individual infestations operate at smaller scales.     

Mastication and prescribed fire impacts on fuels in a 25-year old ponderosa pine plantation, southern Sierra Nevada

Due to increases in tree density and hazardous fuel loading in Sierra Nevadan forests, land management is focusing on fuel reduction treatments to moderate the risk of catastrophic fires. Fuel treatments involving mechanical and prescribed fire methods can reduce surface as well as canopy fuel loads. Mastication is a mechanical method which shreds smaller trees and brush onto the surface fuel layer. Little data exist quantifying masticated fuel beds. Despite the paucity of data on masticated fuels, land managers desire fuel loading, potential fire behavior and fire effects such as tree mortality information for masticated areas. In this study we measured fuel characteristics before and after mastication and mastication plus prescribed burn treatments in a 25-year old ponderosa pine (Pinus ponderosa C. Lawson) plantation. In addition to surface fuel characteristics and tree data collection, bulk density samples were gathered for masticated material. Regressions were created predicting masticated fuel loading from masticated fuel bed depth. Total masticated fuel load prior to fire treatment ranged from 25.9 to 42.9 Mg ha⁻¹, and the bulk density of masticated fuel was 125 kg m⁻³. Mastication treatment alone showed increases in most surface fuel loadings and decreases in canopy fuel loads. Masticated treatment in conjunction with prescribed burning reduced both surface and canopy fuel loads. Detailed information on fuel structure in masticated areas will allow for better predictions of fire behavior and fire effects for fire in masticated fuel types. Understanding potential fire behavior and fire effects associated with masticated fuels will allow managers to make decisions on the possibility of mastication to create fuel breaks or enhance forest health.