Measuring trap efficiency for bark beetles (Col., Scolytidae)

Abstract:  The relative efficiency of cylindrical, linear and cross-barrier traps for trapping bark beetles was investigated based on a theoretical model. Using this model, the effective trap interception area of each trap type was calculated and trap efficiency was defined as the ratio of the effective interception area to the trap surface area. The relative efficiencies of the three trap types were calculated as the ratios of their respective effective interception areas. Based on this approach, assuming random directional movement of dispersing beetles, the order of efficiency of the three trap types, from highest to lowest, was linear, cross-barrier and cylindrical. The expected ratios of trap catches based on the relative efficiencies of the three trap types were fitted to data from trapping experiments with the mountain pine beetle ( Dendroctonus ponderosae Hopkins). In general, there was large variation in trap catches among traps of the same type but the ratios of mean catches per trap conformed to the expected ratios. The results indicate that the model of trap efficiency could be used for designing efficient traps. The methods presented are amenable for assessing the efficiency of other trap designs.     

Biology and host associations of redbay ambrosia beetle (Coleoptera: Curculionidae: Scolytinae), exotic vector of laurel wilt killing redbay trees in the southeastern United States

The redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), and its fungal symbiont, Raffaelea sp., are new introductions to the southeastern United States responsible for the wilt of mature redbay, Persea borbonia (L.) Spreng., trees. In 2006 and 2007, we investigated the seasonal flight activity of X. glabratus, its host associations, and population levels at eight locations in South Carolina and Georgia where infestations ranged from very recent to at least several years old. Adults were active throughout the year with peak activity in early September. Brood development seems to take 50-60 d. Wood infested with beetles and infected with the Raffaelea sp. was similar in attraction to uninfested redbay wood, whereas both were more attractive than a nonhost species. Sassafras, Sassafras albidium (Nutt.) Nees, another species of Lauraceae, was not attractive to X. glabratus and very few beetle entrance holes were found in sassafras wood compared with redbay. Conversely, avocado, Persea americana Mill., was as attractive to X. glabratus as swampbay, P. palustris (Raf.) Sarg., and both were more attractive than the nonhost red maple, Acer rubrum L. However, avocado had relatively few entrance holes in the wood. In 2007, we compared X. glabratus populations in areas where all mature redbay have died to areas where infestations were very active and more recent. Trap catches of X. glabratus and numbers of entrance holes in trap bolts of redbay were correlated with the number of dead trees with leaves attached. Older infestations where mature host trees had been eliminated by the wilt had low numbers of beetles resulting in trap catches ranging from 0.04 to 0.12 beetles per trap per d compared with 4-7 beetles per trap per d in areas with numerous recently dead trees. Our results indicate beetle populations drop dramatically after suitable host material is gone and provide hope that management strategies can be developed to restore redbay trees. The lack of attraction of X. glabratus to sassafras suggests that spread of X. glabratus may slow once it is outside the range of redbay.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

Limonene: attractant kairomone for white pine cone beetles (Coleoptera: Scolytidae) in an eastern white pine seed orchard in western North Carolina

I report on the attraction of the white pine cone beetle, Conophthoru.s coniperda (Schwarz) (Coleoptera: Scolytidae), to traps baited with the host monoterpene limonene in western North Carolina. Both (+)- and (-)-limonene attracted male and female cone beetles to Japanese beetle traps in an eastern white pine, Pinus strobus L., seed orchard near Murphy, NC. Catches of cone beetles were directly proportional to the release rate of (-)-limonene; (+)-limonene was not tested for dose response. Attraction of cone beetles to the pheromone (+/-)-trans-pityol was increased significantly by both enantiomers of limonene. In all experiments, catches of C. coniperda were strongly male biased with no treatment effect on sex ratio. (- )-Limonene had no effect on trap catches of the predator Enoclerus nigripes (Say) to pityol, whereas (+)-limonene interrupted the attraction of E. nigripes to traps baited with pityol. Of six monoterpenes commonly found in white pine cones, only (-)-alpha-pinene elicited attraction of E. nigripes to Japanese beetle traps.     

Impacts of silvicultural thinning treatments on beetle trap captures and tree attacks during low bark beetle populations in ponderosa pine forests of northern Arizona

Our research used a combination of passive traps, funnel traps with lures, baited trees, and surveys of long-term thinning plots to assess the impacts of different levels of stand basal area (BA) on bark beetle tree attack and on trap captures of Ips spp., Dendroctonus spp., and their predators. The study occurred at two sites in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests, from 2004 to 2007 during low bark beetle populations. Residual stand BA ranged from 9.0 to 37.0 m2/ha. More predators and bark beetles were collected in passive traps in stands of lower BA than in stands of higher BA; however, significance varied by species and site, and total number of beetles collected was low. Height of the clear panel passive traps affected trap catches for some species at some sites and years. When pheromone lures were used with funnel traps [Ips pini (Say) lure: lanierone, +03/-97 ipsdienol], we found no significant difference in trap catches among basal area treatments for bark beetles and their predators. Similarly, when trees were baited (Dendroctonus brevicomis LeConte lure: myrcene, exo-brevicomin and frontalin), we found no significant difference for days to first bark beetle attack. Surveys of long-term thinning treatments found evidence of bark beetle attacks only in unthinned plots (approximately 37 m2/ha basal area). We discuss our results in terms of management implications for bark beetle trapping and control.     

Efficacy of traps, lures, and repellents for Xylosandrus compactus (Coleoptera: Curculionidae) and other ambrosia beetles on Coffea arabica plantations and Acacia koa nurseries in Hawaii

The black twig borer, Xylosandrus compactus (Eichhoff) (Coleoptera: Curculionidae: Scolytinae), is a pest of coffee and many endemic Hawaiian plants. Traps baited with chemical attractants commonly are used to capture ambrosia beetles for purposes of monitoring, studying population dynamics, predicting outbreaks, and mass trapping to reduce damage. The objectives of this research were to optimize trapping systems for X. compactus and other ambrosia beetles such as Xylosandrus crassiusculus (Motschulsky) and Xyleborinus saxesenii (Ratzeburg) by comparing efficacy of several attractants, repellents, and trap types. The ability of certain chemicals to act as beetle repellents and thus interfere with trap catch was tested for purposes of protecting host plants from attack. Potential attractants and application methods tested were as follows: ethyl alcohol pouch delivery system, ethyl alcohol vial delivery system, α-pinene in Eppendorf tubes, eugenol bubblecaps, ginger oil bubblecaps, manuka oil bubblecaps, phoebe oil bubblecaps, and an unbaited control. Potential repellents tested were limonene and verbenone. Ethyl alcohol vials were as attractive as ethyl alcohol sleeves, and were more effective than traps baited with eugenol and α-pinene. Japanese beetle traps were more effective for black twig borer trapping than Lindgren funnel traps, and were easier to deploy. Verbenone and limonene significantly reduced trap catch of Xylosandrus compactus and X. crassiusculus, suggesting that they may be effective for reducing attraction to host plants. These results show the importance of developing a combination of several monitoring techniques to enhance management procedures for the black twig borer.     

Influence of environmental and physical factors on capture of Tribolium castaneum (Coleoptera: Tenebrionidae) in a flour mill

Variation in environmental and physical factors within food processing facilities can influence both the distribution of stored-product pests and trapping efficiency. Data from a long-term Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae) monitoring program was used to evaluate spatial variation in captures among trap locations and to determine relationships with environmental and physical variables. From the complete monitoring data set, different subsets were created for the cool and warm seasons, and period of time when environmental and physical factors were measured (2009-2010), with all data sets showing significant differences among trap locations in terms of beetle captures and proportion of time that traps exceeded 2.5 beetles per trap per monitoring period. There was also considerable temporal variation in distribution among the different levels of the mill. Among the environmental and physical variables measured, mean temperature and flour dust accumulation showed the most significant positive relationships with variation of beetle captures at trap locations. More beetles were captured in traps located in close proximity to milling equipment. Presence of equipment near traps was also associated with an increase in flour dust accumulation and temperature. Overall the environmental and physical factors seemed to have a limited influence on variation in captures among trap locations, with temporal variation in distribution perhaps overwhelming potential influences of local trap conditions.     

Mass trapping wild Agriotes obscurus and Agriotes lineatus males with pheromone traps in a permanent grassland population reservoir

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Uraba lugens (Lepidoptera: Nolidae) in New Zealand: pheromone trapping for delimitation and phenology

A synthetic sex pheromone trapping survey of the leaf skeletonizer Uraba lugens Walker (Lepidoptera: Nolidae) demonstrated the unexpectedly widespread distribution of the insect across > 40,000 ha of urban Auckland, New Zealand. A survey of eucalyptus trees planted in parks and other public areas showed a significant spatial correlation between trap catch and breeding populations, validating the trap survey results. Traps in trees showing damage had four-fold higher catches than traps placed in undamaged or nonhost trees, and < 1% of damaged trees with traps failed to catch adult moths. Damage by larval feeding was correlated with male trap catch in the previous generation, offering good prospects for a pest management decision support system, provided that an economic threshold is developed. Catches increased by 3.4-fold in the same georeferenced trapping grid between November and December 2003 and between March and April 2004 across two generations, over the summer. A vertical transect showed that catches increased with height up to the top trap at 13 m (60% of mean tree height). Options for managing the insect will need to overcome the high rate of increase, the rate of spread, and the vertical distribution of the insect on tall eucalyptus trees.     

Can spruce beetle (Dendroctonus rufipennis Kirky) pheromone trap catches or stand conditions predict Engelmann spruce (Picea engelmannii Parry ex Engelm.) tree mortality in Colorado?

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A quarter of a century succession of epigaeic beetle assemblages in remnant habitats in an urbanized matrix (Coleoptera, Carabidae)

We studied the long-term (23-24 years) species turnover and succession of epigaeic beetle assemblages (Coleoptera: Carabidae, incl. Cicindelinae) in three remnant habitats [cottonwood (Populus spp.) and oak (Quercus spp.) stands, and old fields] that are embedded within highly urbanized areas in central Minnesota. A total of 9,710 beetle individuals belonging to 98 species were caught in three sampling years: 1980, 1981 and 2005 in pitfall traps in identical locations within each habitat. Results indicate that there were 2-3 times greater trap catches in 2005 than in 1980 (cottonwood and oak stands, and old fields) and 1.4-1.7 times greater species diversity of beetles in 2005 than in the 1980-1981 suggesting increased habitat association by beetles over time. Although there were no significant differences in catches between 2005 and 1981 (only cottonwood stands and old fields), there was a trend where more beetles were caught in 2005. At the species-level, 10 times more of an open-habitat carabid species, Cyclotrachelus sodalis sodalis LeConte, was caught in 2005 than in 1980. However, trap catches of five other abundant carabid species [Pterostichus novus Straneo, Platynus decentis (Say), Platynus mutus (Say), Calathus gregarius (Say), and Poecilus lucublandus lucublandus (Say)] did not change indicating population stability of some beetle species. These remnant habitats were increasingly colonized by exotic carabid species as Carabus granulatus granulatus Linneaus, Clivina fossor (Linneaus) and Platynus melanarius (Illiger), that were trapped for the first time in 2005. Species composition of epigaeic beetles was quite distinct in 2005 from 1980 with 39 species reported for the first time in 2005, indicating a high turnover of assemblages. At the habitat-level, greatest species diversity was in cottonwood stands and lowest was in old fields, and all habitat types in 2005 diverged from those in 1980s, but not cottonwood stands in 1981. As our sampled areas are among some of the last remnants of the original oak savanna habitats in central Minnesota, we hypothesize that conservation of these sites may be critical to maintaining epigaeic beetle assemblages under increased urbanization pressure.     

Pitfall trap catches of and aphid predation byPterostichus melanarius andPterostichus madidus in insecticide treated and untreated potatoes

Aphids were countedin situ and carabid beetle populations assessed by pitfall trapping betwen 1983–1985 in potato fields in Scotland treated with demeton-S-methyl (DSM) or untreated.Macrosiphum euphorbiae (Thomas) (Homoptera: Aphididae) was the most abundant aphid. Nineteen species of carabid beetles were trapped but the fauna was dominated byPterostichus melanarius (Ill.) andPterostichus madidus (Fabr.) (Coleoptera: Carabidae). Pitfall trap catches of these carabids were variable but generally lower in sprayed than unsprayed plots for a few days after DSM application and higher a week or two later. Of 1800P. melanarius and 910P. madidus dissected, 14.4 per cent and 30.5 per cent respectively, contained aphid remains. The proportion that had eaten aphids often was higher in sprayed than unsprayed plots immediately after treatment. In additional experiments, aphids treated with DSM fell from plants; four to five per cent of the theoretically applied dose of DSM reached the ground in a closed-canopy potato crop; and no mortality occurred whenPterostichus spp. were exposed to DSM in a field bioassay. It is proposed that observed patterns in trap catch ofPterostichus spp. around times of pesticide application may be due in part to aphids falling to the ground after treatment. Beetles that eat these fallen aphids may be less hungry, less active and less likely to be trapped. Later increases in catches may be attributed to hungry, more active beetles, a consequence of prey removal by the insecticide. Although this hypothesis requires further research, it is clear that the effects of insecticide treatment on carabid populations cannot be measured by pitfall trapping alone.     

Trap type, chirality of alpha-pinene, and geographic region affect sampling efficiency of root and lower stem insects in pine.

Root and lower stem insects cause significant damage to conifers, vector phytopathogenic fungi, and can predispose trees to bark beetle attacks. The development of effective sampling techniques is an important component in managing these cryptic insects. We tested the effects of trap type and stereochemistry of alpha-pinene, in combination with ethanol, on catches of the root colonizing weevils (Coleoptera: Curculionidae) Hylobius spp. [mostly Hylobius pales (Herbst)], and Pachylobius picivorus (Germar), the root colonizing bark beetle (Coleoptera: Scolytidae) Hylastes porculus Erickson, and the lower stem colonizing bark beetle Dendroctonus valens (LeConte). We tested for inter-regional differences by conducting similar field assays in the northern (Wisconsin) and southern (Louisiana) United States. The more effective trap type varied with region. Root weevils were caught primarily in pitfall traps in Wisconsin, whereas they were caught mostly in lower stem flight traps in Louisiana. In Wisconsin, root colonizing bark beetles were also caught primarily in pitfall traps, but lower stem colonizing bark beetles were caught primarily in lower stem flight traps. The root feeding weevils preferred (-) over (+)-alpha-pinene in both regions. Some exceptions relating to trap type or gender occurred in southern populations. The two root and lower stem colonizing bark beetles in Wisconsin showed no preference between (+) and (-)-alpha-pinene in combination with ethanol. No bark beetles were caught in the south. Our results suggest that modifying trap type and enantiomeric ratios of monoterpenes for different insect groups and in different regions can improve sampling efficiency for these important pests.     

How varying pest and trap densities affect Tribolium castaneum capture in pheromone traps

The red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), is an important insect pest in food processing facilities. Pheromone trapping is frequently used to monitor red flour beetle populations in structures; however, the optimal trap density and the relationship between trap captures and beetle density is not known. Two experiments were performed concurrently in environmentally controlled 30‐m² walk‐in chambers to determine the relationship between aggregation pheromone trap captures of red flour beetles and beetle and trap number. In one experiment, beetle density was kept constant at 200 individuals per chamber while trap number was varied from 1 to 8, and in the other experiment trap number remained constant at one per chamber while beetle density varied from 20 to 800 individuals. Results indicated that approximately one out of 23 red flour beetles were captured in a trap. Number of beetles captured in traps increased significantly as beetle density increased; however, the proportion of beetles captured remained consistent across beetle densities with a mean of 4.7 ± 0.6% of individuals captured. Trap captures varied significantly with trap placement within experimental chambers, indicating that subtle differences in the trapping environment can influence trap captures. Data suggested that trap densities of 0.07–0.10 m^?2 (2–3 traps per chamber) would maximize trap capture, whereas a trap density of 0.13 m^?2 (four traps per chamber) would maximize the predictive ability of a trapping equation estimating beetle density from trap captures. Results provide information needed to more thoroughly explore how environmental factors might influence red flour beetle trap capture in the absence of changes in beetle density. Further understanding of these relationships will allow for more accurate assessments of absolute beetle density from pheromone trap capture data.     

Phenology and spatial pattern of Typhaea stercorea (Coleoptera: Mycetophagidae) infesting stored grain: estimation by pitfall trapping

The hairy fungus beetle, Typhaea stercorea (L.), occurs frequently in stored grain, often in large numbers. Populations infesting stored barley in Minnesota, corn in South Carolina, and wheat in Florida were sampled by means of grain probe traps. Spatial distribution of the species was examined by contour analysis of trap catch. In South Carolina, corn was sampled at 2 locations over 2 storage seasons, and temperature, moisture content, and malathion residues were measured. These data were used to examine phenology as well as spatial distribution, and showed peak trap catch shortly after harvest in the fall, and in the spring. This pattern followed seasonal changes in grain temperature, but there was no apparent relationship of trap catch to either grain moisture content or malathion residue. The populations of T. stercorea were not distributed randomly, but were largely concentrated in 1 or very few aggregations associated with the "spoutline," a region high in foreign material and broken grain that forms near the center of a bin as it is loaded. However, the spatial patterns were dynamic, even on a very small time scale (week to week). Numbers of insects in aggregations rose and fell, the areas involved expanded and contracted, the centers shifted, and secondary centers appeared and disappeared. These changes were apparently in response to changing patterns of grain temperature and moisture content. Secondary centers of aggregation often formed in warmer grain along bin walls.     

Optimization of a valine:isoleucine methyl ester pheromone blend and comparison of Robbins and Trécé traps for capture of Phyllophaga anxia (Coleoptera: Scarabaeidae) in Rhode Island

Eight ratios of L-valine:L-isoleucine methyl esters were tested in Robbins traps for capture of Phyllophaga anxia (LeConte) adult males. The 90:10, 80:20, and 60:40 ratios of valine:isoleucine were the most effective blends for capture of beetles in Rhode Island. Females were captured in small numbers in some traps but not consistently to any particular blend. Other male Phyllophaga species captured included Phyllophaga fusca (Fr?lich), Phyllophaga forsteri (Burmeister), P. hirsuta (Knoch), and P. marginalis (LeConte). The number of these species collected was low, and it was not possible to determine whether they were attracted to any particular pheromone blend. Peak captures of P. anxia males occurred 31 May in 1999 and 2002 in Kingston, RI. The standard Japanese beetle trap manufactured by Trécé (Adair, OK) captured significantly more beetles than the Robbins trap. Because the Trécé trap is already marketed for Japanese beetles, a lure and trapping system can be adopted for P. anxia.     

Behavioural sequence in the attraction of the bark beetle Ips typographus to pheromone sources

ABSTRACT. The functions of the two synergistic pheromone components, (4S)- cis -verbenol (cV) and 2-methyl-3-buten-2-01 (MB), and the role of ipsdienol in the attraction of Ips typographus (L.) (Scolytidae) to pheromone sources were studied in the field. Absolute and relative beetle catches were compared between several traps placed at and nearby a central pheromone source: a pipe trap containing the source, a surrounding sticky trap, a nearby window trap, and four distant window traps. A higher catch in the outer down-wind distant traps indicated an up-wind anemotaxis to the source. Increased MB release, with cV constant, increased the proportion caught in the central pipe trap, indicating MB as a landing stimulus. Release of MB alone gave a very small catch. Ipsdienol could not substitute for cV in the synergism with MB. An increase of cV, with MB constant, increased the number of beetles caught, but not the proportion caught in the pipe trap. The sex ratio was equal in the window traps, but fell to 30% males in both sticky and pipe traps, showing that a large proportion of the males attracted to the source did not land. The proportion of males in the pipe trap was reduced at the highest cV dose. The results support the idea of each pheromone component having a different relative importance in releasing different steps in the behavioural chain.     

Phenology of Carpophilus spp. (Coleoptera: Nitidulidae) in Stone Fruit Orchards as Determined by Pheromone Trapping: Implications for Prediction of Crop Damage

Traps baited with synthetic aggregation pheromone and fermenting bread dough were used to monitor seasonal incidence and abundance of the ripening fruit pests, Carpophilus hemipterus (L.), C. mutilatus Erichson and C. davidsoni Dobson in stone fruit orchards in the Leeton district of southern New South Wales during five seasons (1991–96). Adult beetles were trapped from September-May, but abundance varied considerably between years with the amount of rainfall in December-January having a major influence on population size and damage potential during the canning peach harvest (late February-March). Below average rainfall in December-January was associated with mean trap catches of<10 beetles/trap/week in low dose pheromone traps during the harvest period in 1991/92 and 1993/94 and no reported damage to ripening fruit. Rainfall in December-January 1992/93 was more than double the average and mean trap catches ranged from 8–27 beetles/week during the harvest period with substantial damage to the peach crop. December-January rainfall was also above average in 1994/95 and 1995/96 and means of 50–300 beetles/trap/week were recorded in high dose pheromone traps during harvest periods. Carpophilus spp. caused economic damage to peach crops in both seasons. These data indicate that it may be possible to predict the likelihood of Carpophilus beetle damage to ripening stone fruit in inland areas of southern Australia, by routine pheromone-based monitoring of beetle populations and summer temperatures and rainfall.     

Effect of Tribolium castaneum (Coleoptera: Tenebrionidae) nutritional environment, sex, and mating status on response to commercial pheromone traps

Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), the red flour beetle, is an important cosmopolitan pest of stored grains. Commercial traps baited with the synthetic aggregation pheromone 4,8-dimethyldecanal (DMD) are used to monitor T. castaneum population densities in storage facilities. However, trap catches may depend on several intrinsic and extrinsic factors. In this study, we explored the effects of beetle nutritional environment, sex, and mating status on the response of T. castaneum to commercial Storgard Dome traps. Beetles raised on a low-nutrition diet were 1.7 times more likely to enter DMD-baited traps compared with beetles that were raised on a high-nutrition diet. Although no sex difference in trap response was found, unmated beetles of both sexes were more responsive to DMD than were mated beetles, and this effect was especially pronounced for beetles reared on a low-nutrition diet. These results suggest that estimating T. castaneum population densities based on trap catches might be improved by incorporating information about the nutritional quality of infested stored products.     

Diurnal flight pattern of Platypus koryoensis (Coleoptera: Platypodinae) in relation to abiotic factors in Korea

The diurnal flight pattern of Platypus koryoensis (Murayama) was examined using sticky traps attached to the trunks of oak trees in central Korea in 2010 and 2011. The flight activities of the beetle were estimated on the basis of 3-h intervals for trap catches from 11:00 to 14:00 on the next day, on June 25–26 and July 1–2, 2010, and on the basis of 2-h intervals for trap catches from 5:00 to 17:00, between June 9 and July 21, 2011 (the peak flight period of the beetle). Over 77% of the beetles were caught from 9:00 to 13:00, with the daily variations in the facing slope. The beetles began to be caught when the air temperature reached around 16 °C, and were the most active when the temperature ranged from 20 to 27 °C. No beetles were caught during rainfall, suggesting that rainfall is one of the factors that hinder beetle flight. The beetles were caught by traps in the east-facing plot earlier than those in the south- and west-facing plots suggesting that the flight behavior of the beetle can be affected by the light. Direction of the beetle flight during the peak of daily flight (from 09:00 to 13:00) was downward along the slope.