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.     

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.     

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.     

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.     

The effects of mechanical fuel reduction treatments on the activity of bark beetles (Coleoptera: Scolytidae) infesting ponderosa pine

Selective logging, fire suppression, forest succession and climatic changes have resulted in high fire hazards over large areas of the western USA. Federal and state hazardous fuel reduction programs have increased accordingly to reduce the risk, extent and severity of these events, particularly in the wildland–urban interface. In this study, we examined the effects of mechanical fuel reduction treatments on the activity of bark beetles in ponderosa pine, Pinus ponderosa Dougl ex. Laws., forests located in Arizona and California, USA. Treatments were applied in both late spring (April–May) and late summer (August–September) and included: (1) thinned biomass chipped and randomly dispersed within each 0.4 ha plot; (2) thinned biomass chipped, randomly dispersed within each plot and raked 2 m from the base of residual trees; (3) thinned biomass lopped-and-scattered (thinned trees cut into 1–2 m lengths) within each plot; (4) an untreated control. The mean percentage of residual trees attacked by bark beetles ranged from 2.0% (untreated control) to 30.2% (plots thinned in spring with all biomass chipped). A three-fold increase in the percentage of trees attacked by bark beetles was observed in chipped versus lopped-and-scattered plots. Bark beetle colonization of residual trees was higher during spring treatments, which corresponded with peak adult beetle flight periods as measured by funnel trap captures. Raking chips away from the base of residual trees did not significantly affect attack rates. Several bark beetle species were present including the roundheaded pine beetle, Dendroctonus adjunctus Blandford (AZ), western pine beetle, D. brevicomis LeConte (AZ and CA), mountain pine beetle, D. ponderosae Hopkins (CA), red turpentine beetle, D. valens LeConte (AZ and CA), Arizona fivespined ips, Ips lecontei Swaine (AZ), California fivespined ips, I. paraconfusus Lanier (CA) and pine engraver, I. pini (Say) (AZ). Dendroctonus valens was the most common bark beetle infesting residual trees. A significant correlation was found between the number of trees chipped per plot and the percentage of residual trees with D. valens attacks. A significantly higher percentage of residual trees was attacked by D. brevicomis in plots that were chipped in spring compared to the untreated control. In lopped-and-scattered treatments, engraver beetles produced substantial broods in logging debris, but few attacks were observed on standing trees. At present, no significant difference in tree mortality exists among treatments. A few trees appeared to have died solely from D. valens attacks, as no other scolytids were observed in the upper bole. In a laboratory study conducted to provide an explanation for the bark beetle responses observed in this study, monoterpene elution rates from chip piles declined sharply over time, but were relatively constant in lopped-and-piled treatments. The quantities of β-pinene, 3-carene, -pinene and myrcene eluting from chips exceeded those from lopped-and-piled slash during each of 15 sample periods. These laboratory results may, in part, explain the bark beetle response observed in chipping treatments. The implications of these results to sustainable forest management are discussed.     

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.     

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.     

The influence of mountain pine beetle outbreaks and drought on severe wildfires in northwestern Colorado and southern Wyoming: A look at the past century

Outbreaks of bark beetles and drought both lead to concerns about increased fire risk, but the relative importance of these two factors is the subject of much debate. We examined how mountain pine beetle (MPB) outbreaks and drought have contributed to the fire regime of lodgepole pine forests in northwestern Colorado and adjacent areas of southern Wyoming over the past century. We used dendroecological methods to reconstruct the pre-fire history of MPB outbreaks in twenty lodgepole pine stands that had burned between 1939 and 2006 and in 20 nearby lodgepole pine stands that were otherwise similar but that had not burned. Our data represent c. 80% of all large fires that had occurred in lodgepole pine forests in this study area over the past century. We also compared Palmer Drought Severity Index (PDSI) and actual evapotranspiration (AET) values between fire years and non-fire years. Burned stands were no more likely to have been affected by outbreak prior to fires than were nearby unburned stands. However, PDSI and AET values were both lower during fire years than during non-fire years. This work indicates that climate has been more important than outbreaks to the fire regime of lodgepole pine forests in this region over the past century. Indeed, we found no detectable increase in the occurrence of high-severity fires following MPB outbreaks. Dry conditions, rather than changes in fuels associated with outbreaks, appear to be most limiting to the occurrence of severe fires in these forests.     

Distribution of saproxylic beetles in a recently burnt landscape of the northern boreal forest of Québec

This study evaluated the importance of burned habitat characteristics as well as the likely dispersal from specific habitats in the distribution of saproxylic beetles the same year as a fire occurred, in burned black spruce stands (Picea mariana [Mill] B.S.P.) in the northern boreal forest of Québec. The distribution of early post-fire saproxylic species was mainly driven by burned habitat attributes at the plot scale (0.04 ha), especially fire severity, suggesting that the effect of environment attributes can act at a relatively fine scale. Some xylophagous and most predaceous species were more abundant in severely burned stands whereas fire severity had the opposite effect on several common mycophagous species. The amount of newly fire-killed trees that could be used as breeding substrates in the burned stands had only a weak positive influence on these functional groups. The great majority of early saproxylic species were weakly associated with the distance from unburned forests or other recently burned patches that could act as potential “source habitats”. Indeed, these variables were of lesser importance than the attributes of the burned habitat. Woody debris that were already present in plots before the fire, potentially serving as local of source-populations for early colonizers, had virtually no influence on the local abundance of species. Many saproxylic species, including some true pyrophilous, clearly showed higher abundance as distance from unburned stands increased. This unexpected relation may reflect that dispersal of insects toward the burnt landscape very shortly after fire could be driven by the higher amount of volatiles released by severely burned forests, which are more likely as distance from unburned forest increased.     

Pinus pinaster Ait. tree mortality following wildfire in Spain

Maritime pine (Pinus pinaster Ait.) is the tree species most affected by wildfire in the Iberian Peninsula. Prediction of the probability of fire-injured tree mortality is critical for management of burned areas, evaluation of the ecological and economic impact of wildfire and prescribed fire planning and application. Pine bark beetles (Scolytidae) frequently attack burned maritime pine stands and cause extensive post-fire mortality throughout the Iberian Peninsula. In the present study, maritime pine trees were monitored for three years following 14 wildfires in four ecotypes in Spain (11 fires in Galicia (Galician ecotype - NW Spain), one fire in Portillo (Meseta-Castellana ecotype - Central Spain), one fire in Rodenal (Rodenal ecotype - Central Spain), and one fire in Genalguacil (Sierra Bermeja ecotype - SW Spain)). Data on tree attributes, crown and bole injury, ground fire severity, Ips sp. presence and tree survival were obtained by examining 3085 trees. Logistic regression models for predicting the probability of delayed maritime pine mortality were developed by use of generalized estimated equations (GEE). An ample range of response to fire damage in mortality was evident among the four ecotypes and different models were fitted for each. The most important variables for predicting tree mortality were total crown volume damaged, presence of Ips sp. attack and cambium kill rating. The results highlight the extensive presence of Ips sp. in burned maritime pine forests and its importance in tree mortality process, the ample range of response of P. pinaster, in terms of post-fire mortality, as well as the need to develop site specific mortality models for the different ecotypes of this species following fire.     

Bark beetles,fuels, fires and implications for forest management in the Intermountain West

Bark beetle-caused tree mortality in conifer forests affects the quantity and quality of forest fuels and has long been assumed to increase fire hazard and potential fire behavior. In reality, bark beetles, and their effects on fuel accumulation, and subsequent fire hazard, are poorly understood. We extensively sampled fuels in three bark beetle-affected Intermountain conifer forests and compared these data to existing research on bark beetle/fuels/fire interactions within the context of the disturbance regime. Data were collected in endemic, epidemic and post-epidemic stands of Douglas-fir, lodgepole pine and Engelmann spruce. From these data, we evaluated the influence of bark beetle-caused tree mortality on various fuels characteristics over the course of a bark beetle rotation. The data showed that changes in fuels over time create periods where the potential for high intensity and/or severe fires increases or decreases. The net result of bark beetle epidemics was a substantial change in species composition and a highly altered fuels complex. Early in epidemics there is a net increase in the amount of fine surface fuels when compared to endemic stands. In post-epidemic stands large, dead, woody fuels, and live surface fuels dominate. We then discuss potential fire behavior in bark beetle-affected conifer fuels based on actual and simulated fuels data. Results indicated that for surface fires both rates of fire spread and fireline intensities were higher in the current epidemic stands than in the endemic stands. Rates of spread and fireline intensities were higher in epidemic stands due, however, to decreased vegetative sheltering and its effect on mid-flame wind speed, rather than changes in fuels. Passive crown fires were more likely in post-epidemic stands, but active crown fires were less likely due to decreased aerial fuel continuity. We also discuss the ecological effects of extreme fire behavior. Information is presented on managing forests to reduce the impact of bark beetle outbreaks and the interplay between management, bark beetle populations, fuels and fire hazard and behavior.     

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.     

Bark beetle (Coleoptera, Scolytidae) outbreak and system of IPM measures in an area affected by intensive forest decline connected with honey fungus (Armillaria sp.)

The character of a bark beetle outbreak, planning system, and implementation of IPM measures in a forest affected by intensive decline connected with honey fungus (Armillaria sp.) is described. It is possible to distinguish two levels of outbreak in the study area. The first level is characterized by spruce mortality connected with yellowing of spruce and presence of plant pathogens, mostly Armillaria sp. Forest stands have disperse infestations of bark beetles. Identification of infested trees and salvage cutting in time are problematic. In case of late processing of infested trees, the second level of outbreak could become manifest. The populations of bark beetle increase. Beetles could attack relatively healthy trees. The classical outbreak with forest edges and spot infestations starts. The identification of infested trees is easier. The application of pheromone trap barriers is effective. Thus, the second level of outbreak could be easily managed. The planning of IPM measures is concerned with localization of various pheromone trap systems in particular forest stands. It is based on field survey, data acquisition, estimation of swarming bark beetle population, and information of pheromone system effectiveness.     

Platypus cylindrus,a vector of Ceratocystis platani in Platanus orientalis stands in Greece

The ambrosia beetle Platypus cylindrus was very common in stands of Platanus orientalis trees in Greece, infesting trees already infected by Ceratocystis platani. The fungus C. platani was isolated from 59% of adult beetles of P. cylindrus collected on the wing in a heavily infected stand of P. orientalis. Transmission of C. platani was demonstrated in 80% of P. orientalis seedlings challenged with P. cylindrus beetles, which were previously fed in cultures of the fungus. In most of the cases, the beetles had bored tunnels in stems, where abundant perithecia of C. platani were observed. Platypus cylindrus adults regularly visited artificially wounded P. orientalis trees in a natural stand. Although this ambrosia beetle normally infests stressed or dead trees, it appears to play a role as a vector of C. platani, especially by visiting wounded trees.     

Post-fire attractiveness of maritime pines (<Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait.) to xylophagous insects

The spectrum and dynamics of xylophagous insects infesting Pinus pinaster after a fire event were studied from November 2007 to December 2008 at Sierra de Outes (Galicia, NW Spain). Insects were collected within and outside the fire area. Relationships among species status, species abundance, tree parameters and tree injury were assessed. Mortality of injured trees was also recorded. Insect colonization built up consistently during the first year after the fire, although the pests did not colonize unburned trees in nearby areas. Thirteen insect species from six xylophagous families were collected. Tomicus spp., Buprestis novenmaculata, Anobium punctatum and Pissodes castaneus were the first species detected. Tomicus spp. (29%) and Ips sexdentatus (23%) were the dominant taxa. Bole char height and soil burn severity were positively correlated with insect presence. Trees with short stem diameter and thinner bark were also preferred. B. novenmaculata presence was positively correlated with crown scorch. Loss of cambium hydration was negatively related to the occurrence of pests, likely because sapwood desiccation and loss of nutrient impede brood development. About 33.3% of the fire-scorched pines died. The results recommend the removal of injured and dying pines after fire, to avoid the rise of pest population threatening recovering trees in the burned areas.     

Ambrosia beetles (Coleoptera: Curculionidae) can directly transmit the fungal pathogens responsible for Rapid ʻŌhiʻa Death

The ecologically and culturally vital tree species, ʻōhiʻa lehua (Metrosideros polymorpha), is threatened by the fungal pathogens Ceratocystis lukuohia and Ceratocystis huliohia, the causal agents of the disease complex called Rapid ʻŌhiʻa Death (ROD). Four invasive ambrosia beetle (Coleoptera: Curculionidae: Scolytinae) species in the Xyleborini tribe colonize ROD Ceratocystis-infested ‘ōhiʻa and produce inoculum through their frass; however, the potential for direct transmission of the ROD fungi by these beetles was unknown. We fulfilled Leach's rules to support insect transmission of ROD by documenting the visitation of these ambrosia beetles to healthy ‘ōhiʻa trees, culturing C. lukuohia and C. huliohia from the ROD-associated beetles using three different collection methods at multiple study sites, and challenging healthy ʻōhiʻa seedlings with beetles that were exposed to both C. lukuohia and C. huliohia cultures. We documented all four invasive ROD-associated ambrosia beetle species including Xyleborinus saxesenii, Xyleborus affinis, Xyleborus ferrugineus, and Xyleborus perforans to regularly visit healthy ʻōhiʻa trees on sticky traps. Viable Ceratocystis propagules were isolated from all species, and C. lukuohia was most commonly isolated of the two ROD-causing fungi. Consistently across all collection techniques, ROD Ceratocystis spp. were detected on just under 3% of all assayed beetles, with the highest detection rate from X. affinis. All four beetle species were capable of directly transmitting both pathogens to healthy ʻōhiʻa seedlings with a high rate of transfer. Ceratocystis spp. are highly virulent pathogens in trees, and a single inoculation can result in tree death, therefore any direct transmission is a cause for concern. After meeting the criteria of Leach's rules, we propose that Xi. saxesenii, X. affinis, X. ferrugineus, and X. perforans are vectors of C. lukuohia and C. huliohia, particularly in areas of high ROD pressure and tree stress.     

Effects of bark girdling on carbohydrate supply and resistance of loblolly pine to southern pine beetle (Dendroctonus frontalis Zimm.) attack

Importance of current photosynthate in the regulation of tree defense against the southern pine beetle (SPB),Dendroctonus frontalis Zimm. (Coleoptera: Scolytidae) was investigated in loblolly pine,Pinus taeda L. Downward translocation of carbohydrate was blocked by removing a ring of bark (girdle) at 3.5 m above ground. Beetle colonization success and tree defensive responses measured as resin flow and induced lesion formation to inoculations with the beetle-associated fungus,Ophiostoma minus (Hedge.) H. and P. Sydow., were assessed above and below girdles. We hypothesized that a decrease in a available carbohydrate would lead to a decrease in tree defense and an increase in SPB colonization success below girdles. At 2 weeks, post-girdling carbohydrates were already reduced by 50% below girdles, but no significant differences in beetle colonization success or defensive responses were measured. At 8 weeks, post-girdling starch was reduced by 93%, sucrose by 44% and resin flow by 44% below girdles. Southern pine beetle adults were more successful in colonizing areas below than above girdles. However, no significant differences were detected in the size of the lesions that formed in response to inoculated fungi. This suggests that the size of the induced lesion may not be a reliable indicator of tree resistance to bark beetle attack as previously believed, and that lesion development is not entirely dependent on available carbohydrate or related to tree defense against beetles. Cambial growth was reduced below girdles with no latewood formed below and 25 rows of latewood formed above girdles. Latewood contains much higher densities of resin ducts than earlywood. This may explain why resin flow was lower and beetles were more successful in colonizing areas of the tree below girdles. Therefore, tree defense against bark beetles may be more complex than a simple supply-and-demand relationship for carbohydrate, and changes in source-sink relationships, as influenced by the environment, may be more important than supply.     

Immediate effects of fire-severity on soil invertebrates in cut and uncut pine forests

We studied how variations in fire severity and the degree of cutting before burning affected soil invertebrates in a Pinus sylvestris forest in central Sweden. A varied depth of burn in the mor layer was obtained by exclusion of rain and addition of fuel in small plots (1 m×2 m) in clear-cut, selectively cut and uncut part of the forest before large-scale prescribed burning took place. Soil samples were taken from the plots immediately before, the day after, and two months after the fire. The overall mortality of invertebrates depended on the proportion of organic soil consumed by the fire, and for individual taxa it ranged between 59 and 100%. Invertebrates that lived deeper in soil suffered lesser mortality than those in the vegetation and litter layers did. Greater mobility in soil (Staphylinidae) or a thick cuticle (Oribatediae, Elateridae) may have contributed to the higher survival observed in these taxa. The beetles Atomaria pulchra (Cryptophagidae), Corticaria rubripes (Lathridiidae), and other fire-favoured insects colonised the burned forest the very day the fire burned. These species preferred the hard-burned plots and the uncut stand for colonisation. Sixty days after the fire, the abundance of invertebrates was lower in the burned cut stands compared to the burned uncut stand. The species composition of beetles in the burned stands was then characterised by a few very abundant fire-favoured species.     

Eight years of seasonal burning and herbicidal brush control influence sapling longleaf pine growth,understory vegetation,and the outcome of an ensuing wildfire

To study how fire or herbicide use influences longleaf pine (Pinus palustris Mill.) overstory and understory vegetation, five treatments were initiated in a 5–6-year-old longleaf pine stand: check, biennial arborescent plant control by directed herbicide application, and biennial burning in March, May, or July. The herbicide or prescribed fire treatments were applied in 1999, 2001, 2003, and 2005. All prescribed fires were intense and averaged 700 kJ/s/m of fire front across all 12 burns. Using pretreatment variables as covariates, longleaf pine survival and volume per hectare were significantly less on the three prescribed fire treatments than on checks. Least-square means in 2006 for survival were 70, 65, 64, 58, and 56% and volume per hectare was 129, 125, 65, 84, and 80 m³/ha on the check, herbicide, March-, May-, and July-burn treatments, respectively. A wildfire in March 2007 disproportionately killed pine trees on the study plots. In October 2007, pine volume per hectare was 85, 111, 68, 98, and 93 m³/ha and survival was 32, 41, 53, 57, and 55% on the check, herbicide, March-, May-, and July-burn treatments, respectively, after dropping trees that died through January 2009 from the database. Understory plant cover was also affected by treatment and the ensuing wildfire. In September 2006, herbaceous plant cover averaged 4% on the two unburned treatments and 42% on the three prescribed fire treatments. Seven months after the wildfire, herbaceous plant cover averaged 42% on the two previously unburned treatments and 50% on the three prescribed fire treatments. Before the wildfire, understory tree cover was significantly greater on checks (15%) than on the other four treatments (1.3%), but understory tree cover was similar across all five treatments 7 months after the wildfire averaging 1.1%. The greater apparent intensity of the wildfire on the previously unburned treatments most likely resulted from a greater accumulation of fuels on the check and herbicide plots that also collectively had a higher caloric content than fuels on the biennially prescribed burned plots. These results showed the destructive force of wildfire to overstory trees in unburned longleaf pine stands while also demonstrating the rejuvenating effects of wildfire within herbaceous plant communities. They caution for careful reintroduction of prescribed fire even if fire was excluded for less than a decade.     

Diversity of Ophiostomatales species associated with conifer-infesting beetles in the Western Carpathians

Bark beetles (Coleoptera: Curculionidae, Scolytinae) are commonly recognised as important agents of tree mortality in coniferous forests of the Western Carpathians. They, together with weevils, are consistently associated with ophiostomatoid fungi. Information regarding conifer beetle-associated fungi in the Western Carpathians remains incomplete and unreliable, particularly with respect to fir-infesting bark beetles. This study aims to clarify associations between fungi in the genera Graphilbum, Leptographium, Ophiostoma and Sporothrix (Ophiostomatales) and their beetle vectors in Norway spruce (Picea abies), European larch (Larix decidua) and silver fir (Abies alba). Samples associated with 20 bark beetle species and weevils were collected from nine stands in Poland and the Czech Republic. Fungi were isolated from adult beetles and galleries. Isolates were identified based on morphology, DNA sequence comparisons for four gene regions (ITS, LSU, ß-tubulin, TEF 1-α) and phylogenetic analyses. In total, 46 distinct taxa were identified, including 25 known and 21 currently unknown species. Several associations between fungi and subcortical insects were recorded for the first time. In addition, O. borealis and O. quercus were detected from A. alba for the first time. The composition of the fungal communities varied among the studied tree species and to a lesser degree among the beetle species. The spruce-infesting bark beetles were commonly associated with species of Leptographium s. l. and Ophiostoma s. str.; the larch-infesting bark beetles were often associated with Ophiostoma s. str. and Sporothrix, while the fir-infesting bark beetles were commonly associated with Ophiostoma s. str. and Graphilbum. The most commonly encountered fungal associates of the examined insects were (a) Grosmannia cucullata, G. piceiperda, Grosmannia sp. 1, Ophiostoma macroclavatum and O. piceae with the spruce-infesting bark beetles; (b) O. pseudocatenulatum and Sporothrix sp. 1 with the larch-infesting bark beetles; and (c) O. piceae, Ophiostoma sp. 2 and Graphilbum sp. 2 with the fir-infesting bark beetles. The differences in fungal associates among the bark beetle species occurring on P. abies, L. decidua and A. alba could be linked to the different habitats that these beetles occupy.