Spatial pattern of pine wilt disease caused byBursaphelenchus xylophilus (Nematoda: Aphelenchoididae) within aPinus thunbergii stand

To understand the mechanism of spread of pine wilt disease caused by the pinewood nematode, Bursaphelenchus xylophilus, which is vectored by a cerambycid, Monochamus alternatus, the spatial distribution of trees weakened by the nematode was examined within a Pinus thunbergii stand from June to October for 4 years. The weakened trees were distributed in a clumped pattern in 1980 and 1981, at an early stage of infestation. In many cases, they showed a double-clumped pattern. The degree of aggregation was higher in June or July than after August. They were uniformly distributed in June or July 1982 and in June 1983 whereas they showed a double-clumped pattern after August. The trees were frequently weakened in June or July when they were near the trees weakened during the previous year. At quadrat sizes of more than 25 m², spatial overlapping was pronounced between trees weakened during June–July of the current year and those weakened in the previous year. The seasonal changes in spatial distribution of weakened trees were explained by the interaction among M. alternatus, B. xylophilus and Pinus trees.     

Oviposition schedules,survivorship curves,and mortality factors within trees of two cerambycid beetles (Coleoptera: Cerambycidae), the Japanese pine sawyer,Monochamus alternatus hope,and sugi bark borer,Semanotus japonicus lacordaire

Summary Oviposition schedules under laboratory conditions, survivorship curves, and mortality factors within trees of two cerambycid beetles, Japanese pine sawyer (JPS),Monochamus alternatus Hope, and sugi bark borer (SBB),Semanotus japonicus Lacordaire, were investigated. Average longevities of reared adults of JPS were 38.1 days for males and 42.3 days for females; those of SBB were 15.7 days for males and 23.8 days for females. It was confirmed that the JPS must feed on the pine branches for full maturation after emergence, but SBB need not; they are able to lay eggs soon after emergence. The average fecundities of JPS and SBB were 32.9 and 90.5, respectively. Thus, the JPS lay fewer eggs for a long time with continuous maturation feeding, whereas the SBB lay more eggs for short time without maturation feeding. Average survivorship curves of JPS within dead pine trees in 8 pine forests were theDeevey's B type, showing a constant mortality through the pre-imaginal stages in the trees. On the other hand those of SBB in 4 cedar stands approached theDeevey's type, suggesting that the high mortality occurred at an early stage in the trees. Average mortalities of JPS between the appearance of oviposition scars and adult emergence in 8 forests ranged from 62.3% to 95.2%. Intraspecific ompetition of JPS resulting from overcrowding in dead pine trees appeared to be main mortality factor. For SBB, as most larvae were killed by resin flow in living trees, this appeared to be the main mortality factor for this species.     

Reducing spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) emergence for hibernation in central British Columbia by felling infested trees

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Population dynamics of Monochamus carolinensis (Col., Cerambycidae) under laboratory conditions

The study of life history dynamics of the pine sawyer beetle Monochamus carolinensis (Olivier) is of interest because the beetles are vectors for the pinewood nematode, Bursaphelenchus xylophilus (Steiner and Buhrer) Nickle, the causal agent of pine wilt in susceptible pine trees. Previous observation of a laboratory colony of M. carolinensis suggested numbers of adult beetles reared from logs varied little in relation to oviposition densities among individual logs. Logs in laboratory colonies are routinely subjected to artificially high oviposition rates. The objectives of this study were to examine the relationship between initial oviposition rate, holding time (time between log cut and oviposition), log volume and area and adult survivorship in laboratory colony. This study examined the number of eggs, larvae, adults and generation survivorship in logs over a 2‐year period. The number of adults emerged per log was not related to the number of eggs laid. Late larval and adult numbers were significantly correlated with log size. Only 12% of the initial cohort completed development and emerged as adults. We attribute the high within‐log mortality of beetles to intraspecific competition and cannibalism.     

Inhibitory Effect of Bursaphelenchus mucronatus (Nematoda: Aphelenchoididae) on B. xylophilus Boarding Adult Monochamus alternatus (Coleoptera: Cerambycidae)

Inhibitory effects of Bursaphelenchus mucronatus on the number of B. xylophilus carried by an adult Monochamus alternatus were investigated using artificial pupal chambers. When pupal chambers were infested with either B. xylophilus or B. mucronatus, the load of B. xylophilus onto the beetle was greater (P < 0.001) than that of B. mucronatus. However, within the pupal chamber there was no difference in the abundance of the third-stage dispersal juveniles, which would molt to the fourth-stage dispersal juveniles to board beetles. The nematode load on beetles that emerged from pupal chambers infested with both Bursaphelenchus species was smaller (P = 0.015) than that of beetles with B. xylophilus alone but greater (P < 0.001) than that of beetles with B. mucronatus alone, suggesting an inhibitory effect of B. mucronatus. As a result of this study, the rate of inhibition of B. mucronatus on molting of third-stage dispersal juveniles of B. xylophilus to fourth-stage dispersal juveniles was 0.65, which resulted in great inhibition on boarding beetles at a rate of 0.7.     

The bimodal adult activity of Monochamus alternatus (Coleoptera: Cerambycidae) caught in pheromone traps in Jeju can be explained by the competitive attractiveness of dying pine trees

The Japanese pine sawyer, Monochamus alternatus Hope (Coleoptera: Cerambycidae), is known to be the primary vector of pinewood nematode Bursaphelenchus xylophilus (Steiner & Buhrer) Nickle that causes pine wilt disease. Adult activity of M. alternatus caught in a pheromone trap on Jeju Island of Korea showed a bimodal form with the first peak in mid to late June and the second peak in mid to late September. The two peaks were separated between mid and late August, showing a valley. Accumulated degree‐days predicted that the emergence of the second generation adults could be possible just before the second peak. But actually no adults of the second generation occurred in the field development experiments of M. alternatus in 2016 and 2017. Pine trees without oleoresin flow (namely dying trees by the infection of pinewood nematode) were abundant during early July to early August. The bimodal adult activity pattern of M. alternatus could be partially explained by the competitive attractiveness of dying trees against pheromone traps, when we accepted the assumption that dying pine trees attract strongly M. alternatus.     

Effects of dung beetles on seedling emergence from herbaceous seeds in the dung of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in a temperate Japanese grassland ecosystem

The effects of dung form and condition and of dung beetles on the emergence of seedlings from herbaceous seeds in sika deer dung were examined in a temperate grassland ecosystem dominated by Zoysia japonica and Hydrocotyle maritima. I conducted field experiments to compare seedling emergence between dung exposed to dung beetles and intact dung using both dung pellets and pats during a typical rainy month (June) and the hottest, drier month (August), when large numbers of seeds of the dominant species were present in the dung. The exposed dung was immediately attacked and broken up by dung beetles, whereas dung protected from the beetles remained intact. In June, at least 12 herbaceous species, including Z. japonica, H. maritima, Mazus pumilus, and Plantago asiatica, emerged from the dung, versus at least six species in August. Decomposition rates of the pellets in June and decomposition scores of the pats in June and August were positively correlated with the number of emerging seedlings, suggesting that the acceleration of decomposition by dung beetles can positively affect seed germination. In this system of interactions among sika deer, herbaceous plants, and dung beetles, sika deer dung prevented seeds from germinating, and beetles had an indirect positive effect on seedling emergence by accelerating decomposition of the dung, although the extent of the effect may depend on the dung type, plant species, and environmental factors.     

Seasonal flight patterns of Ips typographus in southern Sweden and thermal sums required for emergence

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Potential vector switching in the evolution of Bursaphelenchus xylophilus group nematodes (Nematoda: Aphelenchoididae)

To show the importance of vector switching of nematodes in the evolution of the Bursaphelenchus xylophilus group, we tested a hypothesis that “Bursaphelenchus doui (or its ancestor) was transferred by Acalolepta fraudatrix, Acalolepta sejuncta, and/or Monochamus subfasciatus (or their ancestral species) from broad‐leaved trees to conifers, switched vectors from these cerambycid beetles to Monochamus beetles in conifers, and then evolved into the common ancestor of Bursaphelenchus mucronatus and B. xylophilus.” We used a simple nematode‐loading method to beetles and produced 20 binary combinations of five B. xylophilus group species and four cerambycid beetle species in the tribe Lamiini. The affinity of the nematodes for the beetles was examined based on phoretic stage formation of the nematodes. Phoretic stages of B. doui appeared in all beetle species examined, namely Acalolepta luxuriosa, Psacothea hilaris, A. fraudatrix, and Monochamus alternatus, although the affinity of the nematode for M. alternatus was weak. This finding indicates that B. doui could switch vectors to conifer‐using Monochamus beetles after transfer by A. fraudatrix from broad‐leaved trees to conifers. We conclude that vector switching of nematodes could have potentially happened during the evolutionary history of the B. xylophilus group.     

Changes in spatial and temporal distributions of Monochamus beetles along the fire severity in burned Pinus densiflora forests

Wildfire is an important disturbance factor in forest ecosystem and could affect the distribution of insects. Because Monochamus beetles are known vector insects of pine wood nematodes in Korea and burnt pine trees can be used as oviposition habitats for Monochamus beetles, we investigated changes in the spatial (among fire severity classes) and temporal (between years) distributions of M. alternatus and M. saltuarius. From late May 2017 to early October 2018, four multi-funnel traps with pheromone lure were installed in each fire severity class (unburned, light, light-moderate edge, moderate-severe edge, and severe). All traps were replaced fortnightly from May to October for two years. The spatial distribution of the two Monochamus beetles was significantly affected by fire severity. In particular, M. alternatus showed higher catches in the severely burned sites and responded positively to the fire. A similar number of M. saltuarius individuals were observed between 2017 and 2018, while more M. alternatus were caught in 2018 compared to 2017. Consequently, it was confirmed that the dead wood of P. densiflora caused by the wildfire had a great influence on the density of insects, especially M. alternatus.     

A field experiment on dispersal of newly emerged adults ofMonochamus alternatus (Coleoptera: Cerambycidae)

Newly-emerged adults of Monochamus alternatus aged 1 to 5 days were code-numbered with lacquer paint and released by placing them on the trunks of one or two trees in a Pinus thunbergii stand at weekly intervals during the beetle emergence period from 1980 to 1983. Beetles were captured at weekly intervals from one week after the first day of release. Determinations were made on the distance and direction of beetle dispersal during a week after release and analysed by a method of Inoue (1978). When the stand canopy was closed, the rate of beetle's stay on trees was 0.56 per week. The beetles dispersed at random by walk and flight. When the pine stand was sparse, the rate of beetle's stay on trees was 0.02–0.30 per week. They dispersed at random by flight. The average distances traversed were estimated to be 7.1–37.8 m for the first week after emergence. Using other method, the average distance traversed was estimated to be 10–20 m for each week through the first 3 weeks after release. The results of stepwise multiple regression analysis and a simple field experiment suggested that the dispersal of newly-emerged beetles was affected by stand density, number of beetles emerging from individual dead trees, maximum air temperature, and precipitation.     

Ambrosiodmus rubricollis (Eichhoff) (Coleoptera; Curculionidae; Scolytinae) associated with young Tasmanian blue gum trees

Eucalypts are among the most widely planted forest trees in the world, and outside their native Australian range, the main arthropod pests are sap‐sucking insects, defoliators, gall‐making insects and xylophagous beetles. We report on a new association between a polyphagous wood‐boring beetle and Tasmanian blue gum (Eucalyptus globulus Labill.) in Central Portugal. Unidentified wood‐boring insects were found attacking two three‐year‐old E. globulus trees showing signs of decline among otherwise healthy trees in a commercial plantation, in June 2018. Declining trees presented dead twigs and branches, and recently developed epicormic sprouts evident on the trunks. Insects emerging from logs were identified as Ambrosiodmus rubricollis (Eichhoff), a species native to eastern and southern Asia, with the taxonomic identification validated by molecular analysis. To our knowledge, this is the first record of A. rubricollis associated with a eucalypt species worldwide. It is not clear whether the beetles played any significant role on the decline of the trees, but Ambrosiodmus may be potential pests for several tree and shrub species in Europe, as these beetles can transport pathogenic fungi.     

Changes in the content of indole-3-acetic acid in the needles of dwarf pine in the course of year

The changes in the content of indole-3-acetic acid (IAA) were investigated in the youngest needles ofPinus mugo in the course of year. In newly formed needles, the content of IAA increased, starting from July and reaching its maximum in August to October. The IAA content decreased with a decline in the intensity of apical dominance during dormancy but no changes were observed in budbreak (May and June).     

Survival curves of the population ofdendrolimus spectabilis Bulter (Lepidoptera: Lasiocampidae)

Summary Six survival curves were drawn (Fig. 1) from a survey over two generations of the population ofDendro'imus spectabi'is Butler at three different places in the Tokyo University Forest in Aichi during the years from 1957 to 1959. The counting of insect number was done at one month interval except during the winter season. Any apparent mortality expressed as percentage decrease was not related in this survey with the density of insect population at the beginning of each interval. Considering of the magnitude and the variability of each apparent mortality, there were three critical periods of heavy decrease in the generation decline, they were; 1) September to October when newly born larvae were dispersing, 2) December to March when the third instar larvae were overwintering, and 3) August to September for the period of fully grown larvae and pupae.     

Diversity,ecology, and seasonality of sand flies (Diptera: Psychodidae) of the Jenin District (Palestinian Territories)

The diversity, ecology, and seasonality for sand flies from two localities in Jenin District, the Palestinian Territories, were studied. A total of 12,579 sand flies (5,420 Phlebotomus and 7,159 Sergentomyia) were collected during the study period. The genera Phlebotomus and Sergentomyia are represented by 13 and nine species and subspecies, respectively. Species account was given for all collected species. CDC light traps yielded 7,649 (60.8%) of the total captured sand flies, while sticky traps and aspirators contributed to 36.4 and 2.8% of the total collected specimens, respectively. Phlebotomus sergenti and P. syriacus showed two peaks, one in July and one in October. Phlebotomus tobbi showed one peak towards the end of the summer in September and August, while P. papatasi showed a bimodal peaks pattern, one in June and one in October. Phlebotomus canaaniticus showed a peak in August. P. perfiliewi transcaucasicus and P. neglectus showed a peak in October. Sergentomyia dentata showed one peak in August and increasing numbers from June to August, declining afterwards. Other species, such as S. theodori, had one peak in June, S. taizi had steady numbers across the summer, and S. christophersi had a peak in August.     

Stand-level distribution and movement of Platypus quercivorus adults and patterns of incidence of new infestation

Abstract 1 Flying populations of an ambrosia beetle, Platypus quercivorus (Murayama), a vector of an ambrosia fungus Raffaelea quercivora, which causes deciduous oak diebacks in Japan, were determined by sticky screen traps. 2 Platypus quercivorus beetles tended to move upwards along slopes. The highest concentrations of flying beetles usually occured at the upper forest margins. 3 During the period when the number of flying beetles was increasing, the incidence of newly infested trees spread from the epicentre into the forest. During the period when the number of flying beetles was decreasing, the epicentre shrank into the upper forest edge. Newly infested trees did not occur in this period because most trees had already been infested. 4 Near the upper forest edge, where many beetles were highly concentrated throughout the season, the number of new entry holes decreased greatly after the initial attack early in the season, although many adults were present throughout the period.     

Continuous population estimates forDendroctonus frontalis Zimm. (Coleoptera: Scolytidae) occurring in infestations

Infestations of Dendroctonus frontalisZimm. are often observed to enlarge continuously by the colonization of new hosts in a pattern similar to a forest fire. This pattern of infestation growth presents unique problems in quantitatively estimating populations of D. frontalis. Beetle populations on each infested tree in an infestation go through five processes: attack, oviposition, reemergence, survivorship, and emergence. These processes, which have been described mathematically in the literature, each take several days for completion. In order to follow the distribution and abundance of D. frontalis throughout the course of development of a spot, we need a daily estimate of the number of beetles involved in each process on every tree. Since it is not practical to sample each tree daily, we developed a procedure whereby quantitative estimation procedures for within-tree populations were used in combination with the mathematical models for the life processes to produce a daily record of the number of adults successfully attacking trees, the number of eggs oviposited, the number of beetles reemerging, number of beetles surviving within the trees, and the number of beetles emerging. These daily estimates were then summarized for all trees in the spot for the duration of the infestation. The daily record of populations of D. frontalis, used with information on infestation geometry, were suggested to be of value in describing and elucidating several important facets of population dynamics including dispersal patterns within infestations, between tree beetle loss (mortality), and time lags among the various population processes. The information reported can be used to develop simulation models of population dynamics or to validate existing models.     

Synergistic attraction of pine sawyer Monochamus saltuarius (Coleoptera: Cerambycidae) to monochamol and α‐pinene

Monochamus (Coleoptera: Cerambycidae) species are longhorn pine sawyers that serve as insect vectors of the pinewood nematode Bursaphelenchus xylophilus (Nematoda: Parasitaphelenchidae), which are responsible for debilitating pine wilt disease. An aggregation pheromone, 2‐(1‐undecyloxy)‐1‐ethanol (hereafter referred to as monochamol), was shown to be effective at attracting Monochamus species. However, attraction of the pine sawyers to aggregation pheromones varied depending on semiochemicals, including host plant volatiles and kairomones. In this study, we investigated the abilities of monochamol and the host‐plant volatiles α‐pinene and ethanol to attract M. saltuarius in a pine forest in Cheongsong, Gyeongsangbuk‐do, Korea. A total of 91 M. saltuarius (28 males and 63 females) were captured. The combination of monochamol (700 mg) with α‐pinene and ethanol exhibited a synergistic effect on attracting M. saltuarius (11.0 beetles per trap), whereas monochamol alone and a mixture of α‐pinene and ethanol resulted in the capture of 3.2 beetles and 3.6 beetles per trap, respectively. Our results suggest that multi‐funnel traps baited with a blend of monochamol, α‐pinene and ethanol are highly effective for monitoring M. saltuarius and M. alternatus in pine forests.     

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.     

Body size selection inAcanthoscelides alboscutellatus (Coleoptera: Bruchidae)

Summary Direct observations and analyses of selection occurring in natural populations are rare. The biology of the bruchid beetle,Acanthoscelides alboscutellatus, on its host plant,Ludwigia alternifolia, provides an anusual opportunity to study the process of selection on the morphology of an organism under field conditions.A. alboscutellatus larvae mature within the variably dehiscent fruit ofL. alternifolia. At eclosion, adults are confined within indehiscent fruit but are not confined within dehiscent fruit. Beetles can escape from indehiscent fruit only by forcing their bodies through the fruit's apical pore (a circular opening in the top of the fruit). Thus, during the eclosion stage of this beetle's life cycle the relationship between body size and differential fitness appears to be clearly defined. We examined entrapment ofA. alboscutellatus within indehiscentL. alternifolia fruit in a natural population. Only 8.8% of the beetles that attempted to escape were successful. Smaller beetles were trapped within a narrower range of pore diameters than were larger beetles; and trapped beetles had only limited abilities to enlarge fruit pore diameter. These data suggest (1) that escape from indehiscent fruit is regulated by the relationship between adult body diameter and fruit pore diameter and (2) that adult beetles may experience strong selection for small body diameter (size) within idehiscent fruit.