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.     

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.     

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.     

Response of bark beetles and their natural enemies to fire and fire surrogate treatments in mixed-conifer forests in western Montana

Four treatments (control, burn-only, thin-only, and thin-and-burn) were evaluated for their effects on bark beetle-caused mortality in both the short-term (one to four years) and the long-term (seven years) in mixed-conifer forests in western Montana, USA. In addition to assessing bark beetle responses to these treatments, we also measured natural enemy landing rates and resin flow of ponderosa pine (Pinus ponderosa) the season fire treatments were implemented. All bark beetles were present at low population levels (non-outbreak) for the duration of the study. Post-treatment mortality of trees due to bark beetles was lowest in the thin-only and control units and highest in the units receiving burns. Three tree-killing bark beetle species responded positively to fire treatments: Douglas-fir beetle (Dendroctonus pseudotsugae), pine engraver (Ips pini), and western pine beetle (Dendroctonus brevicomis). Red turpentine beetle (Dendroctonus valens) responded positively to fire treatments, but never caused mortality. Three fire damage variables tested (height of crown scorch, percent circumference of the tree bole scorched, or degree of ground char) were significant factors in predicting beetle attack on trees. Douglas-fir beetle and pine engraver responded rapidly to increased availability of resources (fire-damaged trees); however, successful attacks dropped rapidly once these resources were depleted. Movement to green trees by pine engraver was not observed in plots receiving fire treatments, or in thinned plots where slash supported substantial reproduction by this beetle. The fourth tree-killing beetle present at the site, the mountain pine beetle, did not exhibit responses to any treatment. Natural enemies generally arrived at trees the same time as host bark beetles. However, the landing rates of only one, Medetera spp., was affected by treatment. This predator responded positively to thinning treatments. This insect was present in very high numbers indicating a regulatory effect on beetles, at least in the short-term, in thinned stands. Resin flow decreased from June to August. However, resin flow was significantly higher in trees in August than in June in fire treatments. Increased flow in burned trees later in the season did not affect beetle attack success. Overall, responses by beetles to treatments were short-term and limited to fire-damaged trees. Expansions into green trees did not occur. This lack of spread was likely due to a combination of high tree vigor in residual stands and low background populations of bark beetles.     

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.     

Bark beetle-caused mortality in a drought-affected ponderosa pine landscape in Arizona,USA

Extensive ponderosa pine (Pinus ponderosa Dougl. ex Laws.) mortality associated with a widespread severe drought and increased bark beetle (Coleoptera: Curculionidae, Scolytinae) populations occurred in Arizona from 2001 to 2004. A complex of Ips beetles including: the Arizona fivespined ips, Ips lecontei Swaine, the pine engraver beetle, Ips pini (Say), Ips calligraphus (Germar), Ips latidens (LeConte), Ips knausi Swaine and Ips integer (Eichhoff) were the primary bark beetle species associated with ponderosa pine mortality. In this study we examine stand conditions and physiographic factors associated with bark beetle-caused tree mortality in ponderosa pine forests across five National Forests in Arizona. A total of 633 fixed-radius plots were established across five National Forests in Arizona: Apache-Sitgreaves, Coconino, Kaibab, Prescott, and Tonto. Prior to the bark beetle outbreak, plots with mortality had higher tree and stocking compared with plots without pine mortality. Logistic regression modeling found that probability of ponderosa pine mortality caused by bark beetles was positively correlated with tree density and inversely related with elevation and tree diameter. Given the large geographical extent of this study resulting logistic models to estimate the likelihood of bark beetle attack should have wide applicability across similar ponderosa pine forests across the Southwest. This is particularly true of a model driven by tree density and elevation constructed by combining all forests. Tree mortality resulted in significant reductions in basal area, tree density, stand density index, and mean tree diameter for ponderosa pine and for all species combined in these forests. Most of the observed pine mortality was in the 10–35 cm diameter class, which comprise much of the increase in tree density over the past century as a result of fire suppression and grazing practices. Ecological implications of tree mortality are discussed.     

Multiscale spatial variation of the bark beetle Ips sexdentatus damage in a pine plantation forest (Landes de Gascogne,Southwestern France)

Bark beetles are notorious pests of natural and planted forests causing extensive damage. These insects depend on dead or weakened trees but can switch to healthy trees during an outbreak as mass-attacks allow the beetle to overwhelm tree defences. Climatic events like windstorms are known to favour bark beetle outbreaks because they create a large number of breeding sites, i.e., weakened trees and for this reason, windthrown timber is generally preventively harvested and removed. In December 1999, the southwest of France was struck by a devastating windstorm that felled more that 27 million m³of timber. This event offered the opportunity to study large-scale spatial pattern of trees attacked by the bark beetle Ips sexdentatus and its relationship with the spatial location of pine logs that were temporally stored in piles along stand edges during the post-storm process of fallen tree removal. The study was undertaken in a pure maritime pine forest of 1300 ha in 2001 and 2002. We developed a landscape approach based on a GIS and a complete inventory of attacked trees. During this study more than 70% of the investigated stands had at least one tree attacked by I. sexdentatus  . Spatial aggregation prevailed in stands with n≥15$n\ge 15$ attacked trees. Patches of attacked trees were identified using a kernel estimation procedure coupled with randomization tests. Attacked trees formed patches of 500–700 m² on average which displayed a clumped spatial distribution. Log piles stemming from the sanitation removals were mainly distributed along the large access roads and showed an aggregated spatial pattern as well. The spatial relationship between patches of attacked trees and log pile storage areas was analyzed by means of the Ripley’s statistic that revealed a strong association at the scale of the studied forest. Our results indicated that bark beetle attacks were facilitated in the vicinity of areas where pine logs were stored. The spatial extent of this relationship was >1000 m. Similar results were obtained in 2001 and 2002 despite differences in the number and spatial distribution of attacked trees. The presence of a strong “facilitation effect” suggests that log piles should be removed quickly in order to prevent outbreaks of bark beetles.     

Predicting growth and sequestration of carbon by plantations growing in regions of low-rainfall in southern Australia

In regions of Australia of low–medium rainfall (500–800 mm/year), there is growing community and land-owner support for re-planting trees to achieve multiple environmental objectives, particularly amelioration of soil salinity. Sequestration of carbon by newly established trees is not only another important environmental benefit, but also a potential commercial benefit. To obtain estimates of carbon sequestered by species of commercial potential in such regions, we calibrated the carbon (C) accounting model FullCAM to Eucalyptus cladocalyx and Corymbia maculata plantations. This was achieved by harvesting trees of a range in sizes to determine the allometric relationships that most accurately predict biomass and stem density from measures of stem diameter. Predictions of stem diameter were obtained from a forest growth model (3-PG) previously calibrated for these two species. By applying these predictions of changes in stem diameter as the stand matures in our allometric relationships, we estimated changes in partitioning of biomass (between stem, branches, bark, foliage and roots) and stem wood density as the stand matures under scenarios of 500, 600 and 750 mm mean annual rainfall. We found that for both species, regardless of annual rainfall, throughout the rotation 37–50% of carbon sequestered in the total tree biomass was in the stem, 18–27% in both branches and roots, and the remainder in foliage or bark. However, rate of accumulation of carbon was dependent on annual rainfall, with average annual rate of sequestration of carbon in tree biomass and litter during the first rotation of E. cladocalyx (or C. maculata) increasing from 3.68 (or 4.17) to 4.72 (or 4.86) Mg C ha⁻¹ yr⁻¹ as annual rainfall increased from about 500 to 750 mm. Although it was predicted that decomposition negated any accumulation of debris between successive rotations, carbon was predicted to accumulate in sawlog products, given that assumed rates of product decomposition were slightly less than their rate of accumulation. This resulted in a slight increase (<8 Mg C ha⁻¹) in predicted total sequestration of carbon between successive rotations.     

Crown transparency, tree mortality and stem growth of Pinus sylvestris, and colonization of Tomicus piniperda after an outbreak of Gremmeniella abietina

In the year 2000, large areas of forest in Sweden, mainly 30-50 year old Pinus sylvestris (L.) stands, were attacked by the fungus Gremmeniella abietina (Lagerb.) Morelet. The aims of this study were to investigate: (i) the relationship between G. abietina-induced tree crown transparency (CT) and P. sylvestris (L.) tree mortality; (ii) the influence of CT levels on stem growth; (iii) the recovery of the crown; and (iv) the association of CT and colonization by Tomicus piniperda (L.). Thirty-five permanent sample plots were established in five P. sylvestris stands (38-46 years old), infested by G. abietina, and 23 plots in four reference stands, not obviously infested.During the 5 years following the attack, the total mortality amounted to 454 trees ha⁻¹ and 7.8 m² ha⁻¹, on average, in the five infested stands, corresponding to 42% of the trees and 34% of the basal area at the time of the attack. Most of the mortality occurred within 2 years of the attack. The mortality of individual trees (2002-2005) was found to be related to the crown transparency (CT), the position of needle loss within the crown and the tree diameter at breast height. Based on our modeling, the probability of mortality was substantially increased if the initial CT-value was higher than 85%.Growth reductions were detected for individual trees with an initial CT of >c. 40%. In contrast, trees with a low initial CT (<c. 40%) were not affected and even exhibited increased growth. In the five infested stands, the reductions in basal area and volume increment were estimated to be 26-58%, and, 42-73%, respectively, during the five growing seasons after the attacks.The trees in the infested stands that were still alive in spring 2005 had started to recover in terms of CT. Breeding of T. piniperda on the P. sylvestris (L.) stems occurred almost exclusively on stems with a CT > 90%.The data from this study suggest that when a P. sylvestris (L.) stand has been attacked by G. abietina, trees with a CT above 80% should be felled; the remaining trees will have a high probability of survival and resistance to successful breeding by the T. piniperda.     

Predisposition to bark beetle attack by root herbivores and associated pathogens: Roles in forest decline,gap formation,and persistence of endemic bark beetle populations

Bark beetles are largely known for their ability to undergo intermittent population eruptions that transform entire landscapes and pose significant economic hardships. However, most species do not undergo outbreaks, and eruptive species usually exert only minor disturbances. Understanding the dynamics of tree-killing noneruptive species can provide insights into how beetles persist at low densities, and how some spatiotemporal patterns of host predisposition may more likely favor breaching eruptive thresholds than others. Elucidating mechanisms behind low-density populations is challenging, however, due to the requirement of long-term monitoring and high degrees of spatial and temporal covariance. We censused more than 2700 trees annually over 7 years, and at the end of 17 years, in a mature red pine plantation. Trees were measured for the presence of bark beetles and wood borers that breed within the primary stem, root weevils that breed in root collars, and bark beetles that breed in basal stems. We quantify the sequence of events that drive this decline syndrome, with the primary emergent pattern being an interaction between below- and above-ground herbivores and their fungal symbionts. This interaction results in an expanding forest gap, with subsequent colonization by early-successional vegetation. Spatial position strongly affects the likelihood of tree mortality. A red pine is initially very likely to avoid attack by tree-killing Ips beetles, but attack becomes increasingly likely as the belowground complex spreads to neighboring trees and eventually make trees susceptible. This system is largely internally driven, as there are strong gap edge, but not stand-edge, effects. Additional stressors, such as drought, can provide an intermittent source of susceptible trees to Ips beetles, and elevated temperature slightly accentuates this effect. New gaps can arise from such trees as they subsequently become epicenters for the full complex of organisms associated with this decline, but this is not common. As Ips populations rise, there is some element of positive feedback, in that the proportion of killed trees that were not first colonized by root organisms increases. This positive feedback is very weak, however, and we propose the slope between beetle population density and reliance on host stress as a quantitative distinction along a gradient from noneruptive through eruptive species. Almost all trees colonized by Ips were subsequently colonized by wood borers, likely a source of negative feedback. We discuss implications to our overall understanding of cross-scale interactions, between-guild interactions, forest declines, and eruptive thresholds.     

Forest stand dynamics and sudden oak death: Mortality in mixed-evergreen forests dominated by coast live oak

Sudden oak death (SOD), caused by the recently discovered non-native invasive pathogen, Phytophthora ramorum, has already killed tens of thousands of native coast live oak and tanoak trees in California. Little is known of potential short and long term impacts of this novel plant–pathogen interaction on forest structure and composition. Coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica) form mixed-evergreen forests along the northern California coast. This study measured tree mortality over a gradient of disease in three time periods. Direct measurements of current mortality were taken during 2004, representing a point-in-time estimate of present and ongoing mortality. Past stand conditions, c. 1994, were estimated using a stand reconstruction technique. Future stand conditions, c. 2014, were calculated by assuming that, given a lack of host resistance, live trees showing signs of the disease in 2004 would die. Results indicate that coast live oaks died at a rate of 4.4–5.5% year⁻¹ between 1994 and 2004 in highly impacted sites, compared with a background rate of 0.49% year⁻¹, a ten-fold increase in mortality. From 2004 to 2014, mortality rates in the same sites were 0.8–2.6% year⁻¹. Over the entire period, in highly impacted sites, a 59–70% loss of coast live oak basal area was predicted, and coast live oak decreased from 60% to 40% of total stand basal area, while bay laurel increased from 22% to 37%. Future stand structures will likely have greater proportions of bay laurel relative to coast live oak.     

Combining empirical models and the process-based model 3-PG to predict Eucalyptus nitens plantations growth in Spain

Empirical, statistically based models were used to describe the growth and development of Eucalyptus nitens plantations for a range of site productivities and the standard biomass and pulp silvicultural regime currently applied in Northern Spain. The results obtained, along with data gathered from a network of 68 plots, 48 trees felled for biomass estimations and 73 trees sampled for foliar area estimation were used to parameterize the 3-PG model for this species in Northern Spain. Most parameters associated with allometric relationships and partitioning (i.e. bark and branch fraction, basic density, age modifier and mortality) were derived from local data, and the remaining parameters were obtained from published studies on E. nitens or default values previously used for E. globulus. The parameterized model was validated with data from three trials measured from age 3 years until age 8-14 years, and performed better than the empirical model in terms of total stand under bark volume, mean diameter at breast height, basal area and foliar biomass. The process-based model was then used to forecast changes in plantations subjected to a clearwood regime, initializing the model at age 3 years, considering 3 prunings, 2 thinnings and lengthening the rotation to 18 years. This integrated regime was able to provide biomass for bioenergy, pulp or fibreboard wood and also solid wood, with thinning operations assisting the financial viability, and was a potentially good alternative for productive sites.     

Afforestation method affects the isotopic composition of planted Pinus halepensis in a semiarid region of Spain

We used an isotopic approach to evaluate the effects of three afforestation methods on the ecophysiology of an Aleppo pine plantation in semiarid Spain. The site preparation methods tested were excavation of planting holes (H), subsoiling (S), and subsoiling with addition of urban solid refuse to soil (S + USR). Five years after plantation establishment, trees in the S + USR treatment were over three times larger than those in the S treatment, and nearly five-fold larger than those planted in holes. Differences in tree biomass per hectare were even greater due to disparities in initial planting density and pine tree mortality among treatments. Pine trees in the S + USR treatment showed higher foliar P concentration, δ¹³C and δ¹⁵N than those in the S or H treatments. Foliar δ¹⁵N data proved that trees in the S + USR treatment utilized USR as a source of nitrogen. Foliar δ¹³C and δ¹⁸O data suggest that improved nutrient status differentially stimulated photosynthesis over stomatal conductance in the pine trees of the S + USR treatment, thus enhancing water use efficiency and growth. In the spring of 2002, trees in the S + USR treatment exhibited the most negative predawn water potentials of all the treatments, indicating that the rapid early growth induced by USR accelerated the onset of intense intra-specific competition for water. The results of this study have implications for the establishment and management of Aleppo pine plantations on semiarid soils. Planting seedlings at low density and/or early thinning of pine stands are strongly recommended if fast tree growth is to be maintained beyond the first few years after USR addition to soil. Foliar C, O and N isotope measurements can provide much insight into how resource acquisition by trees is affected by afforestation techniques in pine plantations under dry climatic conditions.     

Relative susceptibility of four species of African mahogany to the shoot borer Hypsipyla robusta (Lepidoptera: Pyralidae) in the moist semideciduous forest of Ghana

We examined the relative susceptibility of four mahogany species, Khaya ivorensis, Khaya anthotheca, Entandrophragma angolense, and E. utile, to Hypsipyla robusta attack. Seeds were obtained from one to three parent trees for each species. The research was conducted in the moist semideciduous forest zone in Ghana and used a randomized complete block design. Tree height and diameter and height to first branch were measured until 24 months after out-planting in the field. H. robusta damage was assessed by counting the numbers of shoots attacked, branches, and dead shoots. Khaya spp. grew better but experienced more attack than Entandrophragma spp. The relative susceptibility to H. robusta attack, from most to least, of the four species was: K. anthotheca > K. ivorensis > E. angolense > E. utile. At 24 months, the mean number of shoots attacked per tree ranged from 1.0 for an E. utile seed source to 3.6 on for a K. anthotheca seed source. At 15 months, K. anthotheca and K. ivorensis started branching at about 1.5 m, but height of clear trunk increased over time due to self-pruning. As K. anthotheca grew taller, the number of H. robusta attacks per tree declined. This suggested that selection of genotypes and species that are tolerant of H. robusta attack based on infestation of young plants may not be appropriate. Genetic factors more completely reflecting the response of different species and genotypes to H. robusta attack may manifest themselves at later growth stages.     

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.     

Quantifying spatio-temporal dispersion of bark beetle infestations in epidemic and non-epidemic conditions

Spatio-temporal dispersal of pest species such as bark beetles plays a key role in their population ecology and outbreak dynamics. Understanding the underlying patterns is crucial for applying appropriate management strategies.In contrast to most existing studies which focus on dispersing beetles, we analysed patches of killed trees resulting from bark beetle infestation. The study was based on a 22-year time series of annually captured colour-infrared (CIR) images of the Bavarian Forest National Park (Germany), where Ips typographus L. (Coleoptera, Curculionidae, Scolytinae) propagates undisturbed by human activity. Newly infested patches comprising at least 5 spruce trees were identified in every time step. This investigation of spatio-temporal spread of infestations primarily focused on (i) parameterizing the size and shape of infestation patches, (ii) modelling an infestation gradient and (iii) evaluating the risk of subsequent infestations on landscape scale. We developed a GIS-based distance ring approach to quantify the distance relation of subsequent infestations, including the distribution of potential hosts.Infestation spread was revealed to be strongly distance dependent, following an inverse power law function: on average 65% of new infestations occurred within a 100 m radius of the previous year’s infestations, and 95% within 500 m. ‘Distance’ proved to be a major determinant of I. typographus dispersal on the landscape scale in each time step of the 22-year series we investigated. Infestation distance thus describes the outcome of beetle dispersal very accurately. The time series showed two alternating periods of epidemic and non-epidemic infestation. These gradation stages did not affect the size and shape of infested patches, but epidemics correlated significantly with a higher percentage of infestations within short distances. Additionally, the resulting infestation risk is highly sensitive to the gradation stage, particularly within the first 100 m around source spots where it increases up to 30%.Our study therefore contributes to a better understanding of the outbreak dynamics of I. typographus and suggests concentrating efficient bark beetle management on areas in the close vicinity of previous years’ infestations.     

Tree biomass estimation in regenerating areas of tropical dry vegetation in northeast Brazil

Allometric equations have been developed for various different vegetation types but have rarely been validated in the field and never for dry tropical forest such as caatinga. In three areas of semi-arid Brazil, with regenerating caatinga vegetation, we measured and weighed twelve hundred individuals of four tree species and used the data to validate equations previously determined in mature caatinga. They and several other equations developed for tropical vegetations overestimate the biomass (B) of trees from the regeneration areas by more than 20%, possibly because these trees have reduced crowns, with lower branch masses. We then determined new allometric equations for them, validating equations for one site against data of the others and pooling the data if they were cross-validated. The best equations were power ones, based on diameter at breast height (D), with little improvement by including height, crown area and/or wood density (Caesalpinia pyramidalis, B = 0.3129D^1.8838; Croton sonderianus, B = 0.4171D^1.5601; Mimosa ophthalmocentra, B = 0.4369D^1.8493; and Mimosa tenuiflora, B = 0.3344D^1.9648 and 0.4138D^1.7718).     

Effect of species and spacing of fast-growing nurse trees on growth of an indigenous tree, Hopea odorata Roxb., in northeast Thailand

We tested the effects of species and spacing of nurse trees on the growth of Hopea odorata, a dipterocarp tree indigenous to Southeast Asia, in a two-storied forest management system in northeast Thailand. Eucalyptus camaldulensis, Acacia auriculiformis, and Senna siamea were planted as nurse trees in 1987 at spacings of 4 m × 8 m, 2 m × 8 m, 4 m × 4 m, and 2 m × 4 m in the Sakaerat Silvicultural Research Station of the Royal Forest Department, Thailand. Seedlings of H. odorata were planted in the nurse tree stands at a uniform spacing of 4 m × 4 m and in control plots (no nurse trees) in 1990. Stem numbers of some nurse trees were thinned by half in 1994. The stem diameter and height of all trees were measured annually until 1995 and again in 2007. The mean annual increment (MAI) in volume was estimated as 8.2–10.1 m³ ha⁻¹ year⁻¹ for E. camaldulensis and 0.9–1.2 m³ ha⁻¹ year⁻¹ for S. siamea, smaller than reported elsewhere. This suggests that the site properties were not suitable for them. The MAI of A. auriculiformis was 7.9–9.8 m³ ha⁻¹ year⁻¹, within the reported range. Survival rates of H. odorata in the S. siamea stands and the control plots decreased rapidly during the first 2 years but then stayed constant from 1992. In contrast, survival rates of H. odorata in the E. camaldulensis and A. auriculiformis stands were initially high (>70%), but then decreased after 1995. Stem diameter, tree height, and stand basal area of H. odorata were large in both the S. siamea stands and the control plots from then. The growth of H. odorata was largest in the 2 m × 8 m S. siamea stands. In contrast, it was restricted in the E. camaldulensis and A. auriculiformis stands owing to strong shading by their canopies. Thinning by 50% tended to facilitate the growth of H. odorata temporarily in the E. camaldulensis and A. auriculiformis stands. The stand basal areas of nurse trees and of H. odorata showed a trade-off. These results suggest that the growth of H. odorata was maximized in the S. siamea stands. We assume, however, that the growth of H. odorata could be improved even in the E. camaldulensis and A. auriculiformis stands by frequent or heavy thinning.     

The significance of rotation periods for mycorrhiza formation in Short Rotation Coppice

Rotation periods control not only the above-ground growth but also the assimilate transfer to the root systems in Short Rotation Coppice (SRC). Since assimilates are needed for the nutrient supply of associated mycorrhizal fungi, their control by rotation period length seems most probable. One poplar (Populus nigra × maximowiczii cv. Max 4) and one willow clone (Salix viminalis clone 78–101) cultivated as SRC were investigated on their ectomycorrhiza formation in response to 15 years of continuous different rotation periods (three and six years) at the same test site in Northern Germany. On the poplar clone the frequency of ectomycorrhizae was significantly lower in 6-year than in 3-year rotation. On the willow clone frequency of ectomycorrhizae was not significantly affected, but the portion of dead fine roots was significantly higher in the 6-year than in the 3-year rotation in autumn. In both rotation systems, the frequency of ectomycorrhizal (EM) colonisation was significantly higher in autumn than in spring. Five EM morphotypes were found on the poplar and seven on the willow clone. EM morphotypes which were common on both clones were formed with two fungal partners of the Pezizales (Geopora cervina, Tuber rufum), one of the Agaricales (Laccaria sp.) and one of the Thelephorales (Thelephoraceae). In spring G. cervina constituted the largest part of all observed EM morphotypes on P. nigra × maximowiczii and S. viminalis. The results indicated a selective promotion of EM formation of some Pezizales (Tuber and Peziza spp.) and some Agaricales (Laccaria spp.) due to shorter rotations, and a selective promotion of other Agaricales (Inocybe sp.) and Boletales (Scleroderma spp.) due to longer rotations. This might allow selective manipulation of the mycorrhizal diversity by the selection of the rotation system. A future challenge will be to select which mycorrhizal diversity might be more advantageous for the vitality and biomass production of poplar and willow clones.     

PHENIPS—A comprehensive phenology model of Ips typographus (L.) (Col., Scolytinae) as a tool for hazard rating of bark beetle infestation

We developed the model PHENIPS for spatial and temporal simulation of the seasonal development of Ips typographus at the Kalkalpen National Park in Austria. The model is based on a digital elevation model used for interpolation of temperature and solar radiation to calculate the microclimatic conditions (bark temperature) for the beetles’ development. Additionally, the beetles’ phenology at Kalkalpen National Park was monitored along with air and bark temperature measurements. The onset of host tree infestation in spring was estimated using a lower threshold of 16.5 °C for flight activity and a mean thermal sum of 140 degree-days (dd) from beginning of April 1st onward. Rate of brood development was calculated from accumulated degree-days of hourly temperature data using upper and lower temperature thresholds of 38.9 and 8.3 °C, respectively, and a nonlinear function for calculating effective thermal sums. Re-emergence of parental beetles occurred at a time when 49.7% of the thermal sum for total development (557 dd) was reached. The model includes the discontinuance of the beetle's reproductive activity at a day length <14.5 h. The rate of successful hibernation of established broods is predicted by assessing the developmental stage of initiated generations at the beginning of the cold period. For validation we compared the timing of phenological events in the field with predicted events using both, hourly recorded data at trap trees in the terrain and generated daily topoclimatic data. Using topoclimatic data, the onset of infestation was predicted with a mean absolute error of 1.3 days. The observed onset of emergence of filial beetles in the field was estimated with a mean error of 39 dd. Our PHENIPS explicitly considers the strong effects of regional topography and stand conditions on local air and bark temperature and can be used for precise monitoring of the actual state of bark beetle development at the specific stand/tree level. Using topoclimatic data, PHENIPS simulates the maximum number of generations which is necessary to assess the potential impact of bark beetle outbreaks at regional scale. Further applications of PHENIPS for site-specific hazard rating of bark beetle infestation are discussed.