Individual-based modeling of the spread of pine wilt disease: vector beetle dispersal and the Allee effect

Pine wilt disease is caused by the pinewood nematode Bursaphelenchus xylophilus, which is vectored by the Japanese pine sawyer beetle Monochamus alternatus. Due to their mutualistic relationship, according to which the nematode weakens and makes trees available for beetle reproduction and the beetle in turn carries and transmits the nematode to healthy pine trees, this disease has resulted in severe damage to pine trees in Japan in recent decades. Previous studies have worked on modeling of population dynamics of the vector beetle and the pine tree to explore spatial expansion of the disease using an integro-difference equation with a dispersal kernel that describes beetle mobility over space. In this paper, I revisit these previous models but retaining individuality: by considering mechanistic interactions at the individual level it is shown that the Allee effect, an increasing per-capita growth rate as population abundance increases, can arise in the beetle dynamics because of the necessity for beetles to contact pine trees at least twice to reproduce successfully. The incubation period after which a tree contacted by a first beetle becomes ready for beetle oviposition by later beetles is crucial for the emergence of this Allee effect. It is also shown, however, that the strength of this Allee effect depends strongly on biological mechanistic properties, especially on beetle mobility. Realistic individual-based modeling highlights the importance of how spatial scales are dealt with in mathematical models. The link between mechanistic individual-based modeling and conventional analytical approaches is also discussed.     

Spread of the pinewood nematode vectored by the Japanese pine sawyer: modeling and analytical approaches

The pinewood nematode, Bursaphelenchus xylophilus, is the causative agent of pine wilt of Pinus thunbergii and P. densiflora in Japan. The nematode is vectored by cerambycid beetles of the genus Monochamus. It is inferred to have been introduced from North America early in the 1900s and then to have distributed in China, Korea, and Taiwan. Intensive and/or long-term studies of pine wilt systems have elucidated the pattern and mechanism of the nematode’s spread within a pine stand, dispersal of vector beetles, and spread pattern of pine wilt within a prefecture. The modeling of nematode spread over pine stands, which involves beetle reproduction within a pine stand, has been developing and should elucidate the factors influencing the rate at which the nematode range expands. In this review, we summarize the biologies of the nematode, beetle, and tree, and then characterize the spread of the nematode within a pine stand, locally over pine stands, and regionally over unit administrative districts. Local and regional spreading of the nematode is related primarily to long-distance dispersal by insect vectors and to the artificial transportation of pine logs infested with the nematode and its vector, respectively.     

Effect of Monochamus carolinensis on the Life History of the Pinewood Nematode,Bursaphelenchus xylophilus

The development of Bursaphelenchus xylophilus in pine wood infested with and free of Monochamus carolinensis was investigated. Formation of third-stage dispersal juveniles occurred in the presence and absence of pine sawyer beetles. The proportion of third-stage dispersal juveniles in the total nematode population was negatively correlated with moisture content of the wood. Formation of nematode dauer juveniles was dependent on the presence of the pine sawyer beetle. Dauer juveniles were present in 3 of 315 wood samples taken from non-beetle-infested Scots pine bolts and 81 of 311 samples taken from beetle-infested bolts. Nematode densities were greater in wood samples taken adjacent to insect larvae, pupae, and teneral adults compared with samples taken from areas void of insect activity. Nematodes recovered from beetle larvae, pupae, and teneral adults were mostly fourth-stage dauer juveniles, although some third-stage dispersal juveniles were also recovered. Dauer juvenile density was highest on teneral adult beetles.     

Breach of the northern Rocky Mountain geoclimatic barrier: initiation of range expansion by the mountain pine beetle

Aim Our aim is to examine the historical breach of the geoclimatic barrier of the Rocky Mountains by the mountain pine beetle (Dendroctonus ponderosae Hopkins). This recent range expansion from west of the North American continental divide into the eastern boreal forest threatens to provide a conduit to naïve pine hosts in eastern North America. We examine the initial expansion events and determine potential mechanism(s) of spread by comparing spread patterns in consecutive years to various dispersal hypotheses such as: (1) meso‐scale atmospheric dispersal of insects from source populations south‐west of the Rocky Mountains in British Columbia (i.e. their historical range), (2) anthropogenic transport of infested plant material, and (3) spread of insect populations across adjacent stands via corridors of suitable habitat. Location British Columbia, Canada. Methods We explore potential mechanism(s) of invasion of the mountain pine beetle using spatial point process models for the initial 3 years of landscape‐level data collection, 2004–2006. Specifically, we examine observed patterns of infestation relative to covariates reflecting various dispersal hypotheses. We select the most parsimonious models for each of the initial 3 years of invasion using information criteria statistics. Results The initial range expansion and invasion of the beetle was characterized by aerial deposition along a strong north‐west to south‐east gradient, with additional aerial deposition and localized dispersal from persisting populations in following years. Main conclusions Following deposition of a wave front of mountain pine beetles parallel to the Rocky Mountains via meso‐scale atmospheric dispersal, the areas of highest intensity of infestations advanced up to 25 km north‐east towards jack pine (Pinus banksiana) habitat in a single year. There appeared to be no association between putative anthropogenic movement of infested materials and initial range expansion of the mountain pine beetle across the continental divide.     

The pattern and range of movement of a checkered beetle predator relative to its bark beetle prey

Theoretical studies of predator‐prey population dynamics have increasingly centered on the role of space and the movement of organisms. Yet, empirical studies have been slow to follow suit. Herein, we quantified the long‐range movement of a checkered beetle, Thanasimus dubius, which is an important predator of a pernicious forest pest, the southern pine beetle, Dendroctonus frontalis. Adult checkered beetles were marked and released at five sites and subsequently recaptured at traps baited with pine and pine beetle semiochemicals and located at distances up to 2 km away from the release point. While the pattern of recaptures‐with‐distance at each site provided a modest fit to a simple random‐diffusion model, there was a consistent discrepancy between observed and expected recaptures: a higher than expected proportion of beetles were recaptured at the more distant traps. To account for this deviation, we developed a model of diffusion that allowed for simple heterogeneity in the population of marked beetles; i.e., a slow and fast moving form of the checkered beetle. This model provided a significantly better fit to the data and formed the basis for our estimates of intra‐forest movement. We estimated that on average, one half of the checkered beetles dispersed at least 1.25 km, one third dispersed>2 km, and 5% dispersed>5 km. The source of the heterogeneous dispersal rates were partially due to differences in beetle size: smaller beetles (for both males and females) were more likely to be recaptured away from the release site than larger beetles. The southern pine beetle (prey for the checkered beetle) exhibited no significant heterogeneity in dispersal ability and provided a very good fit to the simple diffusion model. The only difference in dispersal between these two species was that checkered beetles were undergoing greater long‐distance dispersal than the pine beetles (the radius containing 95% of the dispersing individuals was 5.1 km for the checkered beetle and 2.3 km for the pine beetle). Data on the movement of these two species is used to evaluate a general model of spatial pattern formation in a homogeneous environment, and the potential of the checkered beetle as a biological control agent for the southern pine beetle.     

Evolutionary change in a pine wilt system following the invasion of Japan by the pinewood nematode, <Emphasis Type="Italic">Bursaphelenchus</Emphasis><Emphasis Type="Italic">xylophilus</Emphasis>

Pinewood nematode (PWN), Bursaphelenchus xylophilus, is the causative agent of pine wilt disease (PWD) of pine trees and is transmitted by cerambycid beetles belonging to the genus Monochamus. PWN is believed to have been introduced into Japan from North America at the beginning of the 20th century. In this article, we first provide an outline of the PWD system and the range expansion of PWN in Japan and then review the literature, focusing on the virulence of PWN. Virulence is a heritable trait in PWN, with high virulence being closely related to a high rate of reproduction and within-tree dispersal. When two PWN isolates with different virulence levels are inoculated into pine seedlings, the more virulent nematodes always dominate in dead seedlings. In a laboratory setting, many more virulent nematodes board the insect vectors than avirulent ones. The age at which vectors transmit the most abundant PWNs to pine twigs changes during the course of a PWD epidemic. However, the relation between virulence and transmission of PWN remains as yet relatively unknown. Such information would enable ecologists to predict the evolution of the PWD system. In this review we also compare ecological traits between the PWN and the avirulent congener, B. mucronatus.     

Spatial–temporal analysis of species range expansion: the case of the mountain pine beetle, Dendroctonus ponderosae

Aim The spatial extent of western Canada’s current epidemic of mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae, Scolytinae), is increasing. The roles of the various dispersal processes acting as drivers of range expansion are poorly understood for most species. The aim of this paper is to characterize the movement patterns of the mountain pine beetle in areas where range expansion is occurring, in order to describe the fine‐scale spatial dynamics of processes associated with mountain pine beetle range expansion. Location Three regions of Canada’s Rocky Mountains: Kicking Horse Pass, Yellowhead Pass and Pine Pass. Methods Data on locations of mountain pine beetle‐attacked trees of predominantly lodgepole pine (Pinus contorta var. latifolia) were obtained from annual fixed‐wing aircraft surveys of forest health and helicopter‐based GPS surveys of mountain pine beetle‐damaged areas in British Columbia and Alberta. The annual (1999–2005) spatial extents of outbreak ranges were delineated from these data. Spatial analysis was conducted using the spatial–temporal analysis of moving polygons (STAMP), a recently developed pattern‐based approach. Results We found that distant dispersal patterns (spot infestations) were most often associated with marginal increases in the areal size of mountain pine beetle range polygons. When the mountain pine beetle range size increased rapidly relative to the years examined, local dispersal patterns (adjacent infestation) were more common. In Pine Pass, long‐range dispersal (> 2 km) markedly extended the north‐east border of the mountain pine beetle range. In Yellowhead Pass and Kicking Horse Pass, the extension of the range occurred incrementally via ground‐based spread. Main conclusions Dispersal of mountain pine beetle varies with geography as well as with host and beetle population dynamics. Although colonization is mediated by habitat connectivity, during periods of low overall habitat expansion, dispersal to new distant locations is common, whereas during periods of rapid invasion, locally connected spread is the dominant mode of dispersal. The propensity for long‐range transport to establish new beetle populations, and thus to be considered a driver of range expansion, is likely to be determined by regional weather patterns, and influenced by local topography. We conclude that STAMP appears to be a useful approach for examining changes in biogeograpical ranges, with the potential to reveal both fine‐ and large‐scale patterns.     

Transmission of Pinewood Nematode Through Feeding Wounds of Monochamus carolinensis (Coleoptera: Cerambycidae)

Transmission of pinewood nematode, Bursaphelenchus xylophilus, to mature, field grown Scots pines through feeding wounds of Monochamus carolinensis was investigated by caging nematode-infested beetles on pine branches for 24 hours. Nematodes were transmitted to 31 of 64 branches. Frequency of successful transmission was independent of the sex of the beetle but dependent upon beetle age. Transmission frequencies were highest for beetles 2, 3, and 4 weeks after emergence as adults. The number of nematodes transmitted per branch was low and did not differ between beetle sexes or among beetle age categories. The number of nematodes extracted per branch was correlated with the number of nematodes carried per beetle but was not correlated with the feeding area on the branch.     

Effect of the Nematode Contortylenchus brevicomi on Gallery Construction and Fecundity of the Southern Pine Beetle

Field-collected Dendroctonus frontalis were reared in a controlled environment. Male-female beetle pairs retrieved from galleries 1, 2, or 5 wk after introduction into pine bolts were examined for nematode parasites. Data were obtained for each pair on gallery length, egg niche construction, egg viability, and progeny survival. In a separate study, beetle pairs were reared under laboratory conditions for 10 wk. The number of emerged adult progeny of each pair was recorded. Contortylenchus brevicomi, a nematode parasite, was found in 25% of all beetles that established galleries. After 2 and 3 wk, female beetles infected with the nematode had produced fewer eggs and shorter galleries than did uninfected females. Uninfected females mated with nematode-infected males showed similar trends, although the differences in the 2- and 3-wk tests were not significant. Progeny survival or egg viability was not affected by nematode parasitism of either parent beetle. Unikaryon minutum, a microsporidian parasite found in 65% of all colonizing beetles, had no effect on measured variables. The lower fecundity of beetles parasitized by C. brevicomi continued throughout the insect''s reproductive cycle. After 10 wk, nematode-infected beetle pairs produced fewer emerged adult progeny than did uninfected pairs.     

Energy use by the mountain pine beetle (Coleoptera: Curculionidae: Scolytinae) for dispersal by flight

The mountain pine beetle Dendroctonus ponderosae Hopkins is a major native pest of Pinus Linnaeus (Pinaceae) in western North America. Host colonization by the mountain pine beetle is associated with an obligatory dispersal phase, during which beetles fly in search of a suitable host. Mountain pine beetles use stored energy from feeding in the natal habitat to power flight before host colonization and brood production. Lipids fuel mountain pine beetle flight, although it is not known whether other energy sources are also used during flight. In the present study, we compare the level of energy substrates, proteins, carbohydrates and lipids of individual mountain pine beetles flown on flight mills with unflown control beetles. We use a colorimetric method to measure the entire metabolite content of each individual beetle. The present study reveals that mountain pine beetles are composed of more protein and lipid than carbohydrate. Both female and male mountain pine beetles use lipids and carbohydrates as energy sources during flight. There is variation between sexes, however, in the energy substrates used for flight. Male mountain pine beetles use protein, in addition to lipids and carbohydrates, to fuel flight, whereas protein content is not different between flown and control females.     

Dispersal of Monochamus galloprovincialis (Col.: Cerambycidae) as recorded by mark–release–recapture using pheromone traps

The spread of the pine wood nematode (PWN), Bursaphelenchus xylophylus (Nematoda; Aphelenchoididae), the causal agent of the pine wilt disease, is greatly constrained to the dispersal of its vectors, long‐horned beetles of the Monochamus genus. Disease spread at global and regional scales has been mainly caused by human‐mediated transport, yet at a local scale, the short‐ and long‐distance dispersal behaviour of the beetles determine colonization dynamics. Three mark–release–recapture experiments using commercial traps and lures allowed the parameterization of the dispersal kernel under two landscape fragmentation scenarios for the only known European PWN vector, Monochamus galloprovincialis. The respective release of 171 and 353 laboratory‐reared beetles in continuous pine stands in 2009 and 2010 resulted in 36% and 28% recapture rates, yet, at a fragmented landscape in 2011, only 2% of the released 473 individuals could be recaptured. Recaptures occurred as soon as 7–14 days after their release, in agreement with the requirement of sexual maturation to respond to the pheromone–kairomone attractants. Data from the first two experiments were fitted to one mechanical and two empirical dispersal models, from which the distance dispersal kernels could be computed. Derived estimated radii enclosing 50% and 99% of dispersing M. galloprovincialis under continuous pine stands ranged between 250–532 m and 2344–3495 m depending on the replicate and choice of model. Forecasted recaptures in 2011 resulted in a moderate underestimation of long‐distance dispersal, probably influenced by the high degree of habitat fragmentation. In addition, trapping parameters such as the effective sampling area (0.57–0.76 ha) or the seasonal sampling range (426–645 m) could be derived. Observed results, derived dispersal kernels and trapping parameters provide valuable information for the integrated pest management of PWD. Furthermore, estimated dispersal distances indicate that ongoing clear‐cut measures for eradication in the European Union are likely ineffective in stopping the vectors dispersal.     

The red turpentine beetle, <Emphasis Type="Italic">Dendroctonus valens</Emphasis> LeConte (Scolytidae): an exotic invasive pest of pine in China

An exotic invasive pest of pines, the red turpentine beetle, Dendroctonus valens LeConte (Scolytidae) (RTB), was first detected in Shanxi Province, northern China, in 1998 and started causing widespread tree mortality there in 1999. This outbreak continues and has spread to three adjacent provinces, causing unprecedented tree mortality. Although it is considered a minor pest of pines in North America, RTB has proven to be an aggressive and destructive pest of Pinus tabuliformis, China’s most widely planted pine species. The bionomics and occurrence, distribution, response to host volatiles, and host preference of this pine beetle in China are compared with what is known of the beetle in its native range in North America. Factors likely contributing to D. valens success in China and control of the beetle outbreak are discussed. (−)-β-pinene was shown to be the most attractive host volatile for D. valens from the Sierra Nevada of California, whereas 3-(+)-carene is the most attractive host volatile for beetles in China. Monocultures of Pinus tabuliformis, several consecutive years of drought conditions and warm winters have apparently factored D. valens invasion and establishment in China.     

Parasites and Dung Beetles as Ecosystem Engineers in a Forest Ecosystem

Dung beetles serve as the intermediate host for Streptopharagus pigmentatus, a nematode parasite that infects an old world primate, the Japanese Macaque (Macaca fuscata). This study compares the behaviors of infected and uninfected beetles in both transmission dynamics and the ecological role of the parasite. The results suggest that parasitism does not alter the beetle’s use of shelter or choice of substrate on Yakushima Island, Japan. However, infected beetles consume significantly less feces. Dung beetles remove the majority of fecal material in this forest ecosystem, eliminating breeding grounds for many insect pests and burying nutrients that are essential for plant health. Thus, the nematode parasite S. pigmentatus, by altering its host’s behavior, changes the availability of fecal resources to both plant and animal communities and should therefore be classified as an ecosystem engineer.     

Application of harmonic radar to analyze dispersal behavior of the Japanese pine sawyer beetle,Monochamus alternatus (Coleoptera: Cerambycidae)

Monochamus alternatus (Hope) is a severe wood‐boring pest in coniferous forests and a major vector of pine wilt disease in East Asia. Harmonic radar is a powerful tool for studying the dispersal behavior of insects and it could be applied to control pine wilt disease. In this study, we validated the application of harmonic radar for analyzing the dispersal behavior of M. alternatus beetles in a field environment. We determined the wing capacities of the beetles and the effects of electronic tagging and marking on their movement, flight ability, survivorship, and food consumption in the laboratory to confirm the suitability of this technique. The detection rate and recovery rate for beetles were analyzed separately using radar on caged pine stands and in the field environment. The results showed that the minimum wing capacity of the Japanese pine sawyer was 24.9 mg, which was seven times the weight of the electronic tag (3.5 mg). Marking and tagging the beetles had no significant adverse effects on their movement, flight capacity, food consumption, and survivorship. The detection rate using the radar system and recovery rate based on visual observations of the beetles in caged pines were both 95.6%. However, in the field environment, the detection and recovery rates were only 55.6% and 37.8% after one week, respectively, and 33.3% and 7.8% after two weeks. Harmonic radar is a promising technique for studying the dispersal behavior of the Japanese pine sawyer, but its performance is not satisfactory and major improvements are required for both the radar system and electronic tags.     

Yearly changes in dispersal and life-history traits of Monochamus alternatus Hope with reference to its outbreak

The Japanese pine sawyer, Monochamus alternatus Hope, is the primary vector of the pinewood nematode, Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle, the causative agent of pine wilt disease in East Asia. The range of B. xylophilus expands through the dispersal capability of its vectors and transport of host trees infested with the pathogenic nematode and its vector. Outbreaks of M. alternatus populations occur together with the epidemics of pine wilt disease, because the insect reproduces on host trees recently killed by the disease. We measured some dispersal and life-history traits of adults for four years to determine the change in flight capability and life history of a field population of beetles in relation to an outbreak. The population monitored exhibited an outbreak and subsequent collapse. The greatest mean body mass, largest area of hind wings, smallest wing load, and shortest preoviposition period were observed in the year of outbreak. By contrast, there was no difference in the ovariole number between pre-outbreak (latent) and outbreak years. The greatest mean hind wing area and smallest wing load suggest likely result in greater flight performance. As other studies showed, adult body mass is related positively to the flight performance and oviposition rate. Moreover, a shortened preoviposition period leads to a high reproduction rate. Thus, adults in outbreak populations are “superdispersers” because they are likely to have enhanced flight capability and reproduction power. This suggests that M. alternatus populations at the onset of a population outbreak enhance the expansion rate of B. xylophilus range more than those during the latent and pre-outbreak periods.     

Walking while Parasitized: Effects of a Naturally-Occurring Nematode on Locomotor Activity of Horned Passalus Beetles

Internal parasites typically are associated with a range of negative effects on their hosts, including reduced energy, which can manifest in behavioral alterations. With this in mind, we examined effects of a naturally-occurring nematode parasite, Chondronema passali, on locomotor activity level in horned passalus beetles, Odontotaenius disjunctus from Georgia, USA. This parasite is not well-studied but can number in the thousands in severely parasitized hosts. Prior study in our lab revealed that parasitized beetles actually consume more wood than unparasitized ones do, leading us to ask here, if parasitized beetles are also more physically active. Beetles were collected from nearby forests and housed individually in our lab. We created a simple tabletop arena to observe beetle locomotor activity, which was gridded and included small stones and paper objects. We allowed individual beetles to traverse the arena for 5 min and recorded the number of grid squares crossed. Then, beetles were dissected to determine parasite presence and level of infection (on a categorical scale). A total of 140 beetles were examined across three collections. Statistical analyses of locomotor activity revealed parasite severity predicted locomotor activity, but paradoxically, lightly-infected beetles were twice as active as those without this nematode. Activity diminished with increasing worm burdens thereafter, but even the group with the most severe burdens did not move less than those with no worms. From these results we conclude that this parasite does not result in overall reduction in activity, but rather it appears to come with heightened locomotion. Alternatively, this result could stem from the fact that more active beetles are simply more likely to contract the parasite.

     

Newly Discovered Transmission Pathway of Bursaphelenchus xylophilus from Males of the Beetle Monochamus alternatus to Pinus densiflora Trees via Oviposition Wounds

The transmission of Bursaphelenchus xylophilus from Monochamus alternatus males to Pinus densiflora trees via oviposition wounds has been determined. Nematode-infested males, with mandibles fixed experimentally to prevent feeding, were placed for 48 hours with pine bolts containing oviposition wounds that had been made by nematode-free females. After removal of the nematode-infested males, the pine bolts were held for 1 month and then examined for the presence of nematodes. Reproducing nematode populations were recovered from pine bolts that were exposed to male beetles carrying a high number of nematodes. No reproducing nematode population could be recovered from pine bolts exposed to beetles with a small number of nematodes. Nematode reproduction in the pine bolts was not related to the number of oviposition wounds per bolt. Fourth-stage dispersal B. xylophilus juveniles, collected from beetle body surfaces, were inoculated on pine bolt bark 0, 5, 10, and 15 cm away from a single artificial, small hole. These dauer juveniles successfully entered some bolts. The probability of successful nematode reproduction decreased with increased distance between inoculation point and artificial hole. The results indicated that B. xylophilus can move a significant distance to oviposition wounds along the bark surface and enter a tree via the wounds. The new transmission pathway is considered important for the nematode to persist in pine forests such as in North America where pine wilt disease does not occur.     

Dispersal patterns of exotic forest pests in South Korea

Abstract Invasive species have potentially devastating effects on ecological communities and ecosystems. To understand the invasion process of exotic forest pests in South Korea, we reviewed four major species of exotic forest pests: the pine needle gall midge (Thecodiplosis japonensis), pine wilt disease caused by the pine wood nematode (Bursaphelenchus xylophilus), the fall webworm (Hyphantria cunea) and the black pine bast scale (Matsucoccus thunbergianae). We consider their biology, ecology, invasion history, dispersal patterns and related traits, and management as exotic species. Among these species, the dispersal process of fall webworm was linear, showing a constant range expansion as a function of time, whereas the other three species showed biphasic patterns, rapidly increasing dispersal speed after slow dispersal at the early invasion stage. Moreover, human activities accelerated their expansion, suggesting that prevention of the artificial movement of damaged trees would be useful to slow expansion of exotic species. We believe that this information would be useful to establish management strategies for invasion species.     

Long‐distance dispersal of non‐native pine bark beetles from host resources

1. Dispersal and host detection are behaviours promoting the spread of invading populations in a landscape matrix. In fragmented landscapes, the spatial arrangement of habitat structure affects the dispersal success of organisms. 2. The aim of the present study was to determine the long distance dispersal capabilities of two non‐native pine bark beetles (Hylurgus ligniperda and Hylastes ater) in a modified and fragmented landscape with non‐native pine trees. The role of pine density in relation to the abundance of dispersing beetles was also investigated. 3. This study took place in the Southern Alps, New Zealand. A network of insect panel traps was installed in remote valleys at known distances from pine resources (plantations or windbreaks). Beetle abundance was compared with spatially weighted estimates of nearby pine plantations and pine windbreaks. 4. Both beetles were found ≥25 km from the nearest host patch, indicating strong dispersal and host detection capabilities. Small pine patches appear to serve as stepping stones, promoting spread through the landscape. Hylurgus ligniperda (F.) abundance had a strong inverse association with pine plantations and windbreaks, whereas H. ater abundance was not correlated with distance to pine plantations but positively correlated with distance to pine windbreaks, probably reflecting differences in biology and niche preferences. Host availability and dispersed beetle abundance are the proposed limiting factors impeding the spread of these beetles. 5. These mechanistic insights into the spread and persistence of H. ater and H. ligniperda in a fragmented landscape provide ecologists and land managers with a better understanding of factors leading to successful invasion events, particularly in relation to the importance of long‐distance dispersal ability and the distribution and size of host patches.     

Tree response and mountain pine beetle attack preference,reproduction and emergence timing in mixed whitebark and lodgepole pine stands

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