Restricted dispersal in a flying beetle assessed by telemetry

Many insects living in ancient trees are assumed to be threatened as a result of habitat loss and fragmentation. It is generally expected that species in habitats with low temporal variability in carrying capacity have lower degree of dispersal in comparison to those in more ephemeral habitats. As hollow trees are long-lived, species in that habitat are expected to be sensitive to habitat fragmentation, due to a low capacity to establish new populations far from present ones. Using radio telemetry, we studied the dispersal for a beetle, Osmoderma eremita, living in hollow trees. O. eremita exhibited philopatry and only dispersed over short ranges. About 82–88% of the adults remained in the tree where they were caught. All observed dispersal movements ended up in nearby hollow trees and 62% in the neighbouring hollow tree. These results corroborate the suggestion that habitat fragmentation may be detrimental to insects living in temporally stable but spatially variable habitats. In order to preserve such species, we propose that conservation efforts should be focused on maintaining or increasing the number of suitable trees in and near presently occupied stands.     

Geographic variation in otolith morphology among freshwater populations of Aphanius dispar (Teleostei,Cyprinodontiformes) from the southeastern Arabian Peninsula

Aphanius Nardo is a large genus of teleost fishes in the Old World, with 19 described species. Several of these species have only recently been recognized and additional species can be expected from isolated populations in remote areas. We show here that otolith morphology and statistical analyses of otolith variables can contribute to the detection of genetic differentiation in Aphanius. We studied samples of eight Aphanius dispar populations from the southeastern part of the Arabian Peninsula. Two populations originate from freshwater habitats far inland and probably have been isolated since the late Holocene some 4,000 years ago, three populations come from freshwater habitats with occasional connections to the sea, one population originates from a coastal site, and two were artificially introduced populations. The coastal population is interpreted as possessing the basic otolith type of A. dispar. The basic otolith type also occurs in fishes from the freshwater habitat which is located closest to the coast. Otoliths from the two other freshwater populations with occasional connections to the coast differ slightly from the basic type. However, the two populations from the long‐term isolated freshwater habitats far inland show distinct morphological changes. Our results are consistent with the hypotheses that i) otolith morphology is primarily genetically determined and is little influenced by physical parameters of the habitat, and ii) isolated A. dispar populations may be capable of evolving into new species within short periods of time. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Isolation from forest habitats reduces chances of the presence of <Emphasis Type="Italic">Osmoderma eremita</Emphasis> sensu lato (Coleoptera,Scarabaeidae) in rural avenues

Decline and fragmentation of natural habitats, such as old-growth forests, reduces their availability in the landscape. The solution to this problem for many forest-dwelling species, may be colonization of alternative habitats, such as parks, orchards or rural avenues, located in the highly fragmented agricultural landscape. Our main objective was to determine the effect of both habitat quality parameters and isolation from potential forest habitats, as primary habitats, on the occurrence of the hermit beetle (Osmoderma) in rural avenues in south-western Poland. The study was based on the results of an inventory of the species in 201 rural avenues within an area of approx. 30,000 km². Occurrence of the hermit beetle in such alternative habitats was affected by both habitat quality parameters and connectivity with suitable forest habitats. The species occurrence in an avenue was significantly positively affected by mean tree diameter and diversity of tree species, but probability of occurrence decreased as isolation of avenue from the deciduous forest increased. Moreover, in the study area the hermit beetle seemed to avoid alleys with a large proportion of Acer platanoides, Fraxinus excelsior and Populus spp. Spatially isolated roadside avenues have limited value in the preservation of the hermit beetle in the long term conservation management of the species. Conservation plans in such habitats should therefore take into account surrounding suitable habitats.     

Control of invasive species for the conservation of biodiversity in Mediterranean islands. The LIFE PonDerat project in the Pontine Archipelago,Italy

We present the EU LIFE PonDerat project, which is aimed at restoring the natural ecosystems of the Pontine Archipelago, a group of islands located off the western coast of Italy. The spread of invasive species is a major environmental threat on these islands, which are rich in rare habitats and endemic species and are important sites for the conservation of Mediterranean biodiversity. The project focuses on the conservation of species and habitats that are protected by EU laws but are currently threatened by introduced plants and animals. The main targets of the control measures are black rats, feral goats, mouflons and invasive plants of the genus Carpobrotus. Conservation measures focus on the shearwaters Puffinus yelkouan and Calonectris diomedea, which are endemic to the Mediterranean and are listed in Annex I of the European Bird Directive. Conservation measures also focus on island habitats of great biogeographical value, which are listed in Annex I of the European Habitats Directive and are seriously threatened by introduced herbivores and by invasive plants. The main outcome expected from this project is that the restoration of ecosystem functions will enhance the breeding success and population size of shearwaters and lead to the recovery of target habitats.     

Osmoderma eremita as an indicator of species richness of beetles in tree hollows

The beetle Osmoderma eremita has received much attention in the last few years, as it is among those species with the highest priority in the European Union's Habitat Directive. In this paper the species is evaluated as a potential indicator and umbrella species for the endangered beetle fauna in tree hollows. To be useful as an indicator of a species-rich fauna it should be easy to inventory and be strongly correlated with the presence of other species. An umbrella species is a species which is so demanding that the protection of this species will automatically save many others. The species richness of saproxylic beetles and occupancy of O. eremita were surveyed in tree hollows in an area in southeastern Sweden by assessing the presence/absence of living and dead adults (including fragments) and larvae. The species richness was higher when O. eremita was present, both at tree and stand level. Several threatened species were associated with the presence of O. eremita, whereas others did not correlate with the occurrence of O. eremita. As O. eremita is easy to find and identify, it is useful as an indicator of stands with a rich beetle fauna in tree hollows. Osmoderma eremita can be used as an umbrella species, because if measures are taken to conserve O. eremita, many other species in the same habitat are also conserved. However, there are some beetles in tree hollows which seem to be more sensitive to habitat fragmentation than O. eremita, and may go extinct if only O. eremita is taken into consideration.     

Beetles succession on different microhabitats of small mammals in Riyadh,Kingdom of Saudi Arabia

The decay of small carcasses is considerably neglected although it may provide important forensic data. This study differentiates the beetle species correlated with rat, Rattus norvegicus and mouse, Mus musculus (L. 1758) carcasses placed in two different habitats at King Saud University in western Riyadh. Decay processes and beetle succession were analyzed from January to February 2018. We collected an average of 268.67 beetles belonging to nine species from eight families, including Histeridae, Dermestidae, Cleridae, Tenebrionidae, Staphylinidae, Coccinellidae, Melolonthinae, and Pyrrhocoridae. Collected specimens were identified morphologically and by molecular techniques based on the mitochondrial cytochrome oxidase subunit I (COI) gene. Their presence rose in the bloated and decaying stages of decomposition. The beetle communities differed between the two habitats. In addition, a distinctive beetle community was found in the decaying stage compared to the other stages. The lowest number of beetles was reported in the dry stage. The abundance of beetles was associated with carcass size (rat or mouse) and habitat type.     

Offspring number in a livebearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae): reduced fecundity and reduced plasticity in a population of cave mollies

Life history traits within species often vary among different habitats. We measured female fecundity in mollies (Poecilia mexicana) from a H₂S-rich cave and from a neighbouring surface habitat, as well as in laboratory-reared individuals of both populations raised in either light or continuous darkness. Compared to conspecifics from surface habitats, cave-dwelling P. mexicana had reduced fecundity (adjusted for size) in the field. In the laboratory, the fecundity of surface mollies was higher in light than in darkness, whereas fecundity in the cave mollies was almost unaffected by the ambient light conditions. Our results suggest a heritable component to the reduction in fecundity in female cave mollies. Moreover, the reduced plasticity in fecundity of cave mollies in response to light conditions might be an example of genetic assimilation or channelling of a life history trait in a population invading a new environment.     

Landscape effects of transgenic cotton on non-target ants and beetles

Transgenic crops producing toxins from Bacillus thuringiensis (Bt) can be planted in the same field for many years, and many insects exploiting such crops must disperse to other habitats to persist. Accordingly, effects of transgenic crop farming could accumulate through time and affect insect populations across agricultural landscapes. We monitored the population density of seven ant genera and beetle families and of rare ants and beetles in 84 non-cultivated sites abutting agricultural fields in Central Arizona. We assessed the short-term (during planting year) and long-term (over 5–6 years) landscape effect of farming Cry1Ac cotton on ant and beetle density in non-cultivated sites, in addition to several local and regional variables. Landscape variables (e.g., sequence of crops planted in neighbouring fields, crop diversity, and abundance) were more frequently associated with insect density than local variables (e.g., plant productivity and diversity in non-cultivated sites). In the short-term, use of Bt relative to non-Bt cotton in neighbouring fields was positively associated with density of one ant and two beetle groups in non-cultivated sites. However, acreage of Bt cotton located within 1 km from non-cultivated sites had more negative effects than acreage of non-Bt cotton on density of one ant and one beetle group. In the long-term, the proportion of years that Bt cotton was planted in neighbouring fields was positively associated with ant density but not beetle density. Results suggest that the farming of Bt cotton in neighbouring fields frequently resulted in positive short- and long-term landscape effects on ants and beetles in non-cultivated sites, while Bt cotton planted farther away had less frequent negative short-term impacts.     

A new cave-dwelling and endemic species of the genus Pholeuonopsis (Coleoptera, Leiodidae) from Serbia

In the present study, a new species of a cavernicolous beetle is described from Serbia: Pholeuonopsis (Pholeuonopsis) zlatiborensis sp. n. (Leptodirini, Cholevinae, Leiodidae). All important taxonomic features have been thoroughly analyzed and illustrated. The new species probably belongs to an old phyletic lineage of mesogeid origin, like other known Pholeuonopsis species inhabiting cave and endogean habitats in the western part of the Balkan Peninsula. Therefore, the species is both relict and endemic inhabitant of cave habitats in Serbia.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

河西走廊天然固沙植被区地表甲虫多样性及其对沙漠化的指示作用

干旱、半干旱区沙漠化强烈影响动植物分布及多样性,地表甲虫是荒漠中主要的动物类群,它们对沙漠化引起的植被和土壤环境变化响应十分敏感。鉴于此,以河西走廊中部张掖绿洲外围的天然固沙植被区作为研究区,依据沙漠化发育程度选择流动沙丘（ASD）、丘间低地（IL）、半固定（SFSD）和固定沙丘（FSD）4种生境,调查了地表甲虫群落组成及影响甲虫分布的植被和土壤环境。研究发现,4种生境地表甲虫群落组成明显不同并存在季节变异,5月ASD与IL、SFSD和FSD生境地表甲虫群落的相异性大于8月。5月和8月SFSD生境地表甲虫活动密度均显著高于其他生境,8月FSD生境地表甲虫多样性指数显著高于其他生境。不同大小甲虫对沙漠化的响应模式不同,大中型甲虫对沙漠化的响应较小型甲虫敏感,这在5月表现尤为明显。地表甲虫与环境因子的RDA分析结果表明,12个植被和土壤环境因子解释了49.8%的地表甲虫群落变异,其中植被环境解释了甲虫群落变异的16.3%,土壤环境解释了甲虫群落变异的4.2%,植被和土壤环境相互作用解释了甲虫群落变异的29.3%。pRDA分析结果表明,草本物种丰富度、灌木盖度、土壤有机碳含量和粗砂含量是影响地表甲虫分布的主要环境因子,它们解释了43.7%的地表甲虫群落变异。Pearson相关分析表明,草本物种丰富度与地表甲虫活动密度呈显著正相关,而与地表甲虫均匀度呈显著负相关;灌木盖度与地表甲虫多样性呈显著正相关;地表甲虫物种丰富度与灌木盖度和草本物种丰富度均呈显著正相关。此外,研究还发现戈壁琵甲、克氏扁漠甲、中华砚甲和甘肃齿足象可以用于指示FSD生境,东鳖甲属昆虫可以用于指数SFSD生境,谢氏宽漠王可以用于指示IL及ASD生境。     

Modeling the effects of anthropogenic exploitation and climate change on an endemic stag beetle,Lucanus miwai (Lucanidae), of Taiwan

Loss of biodiversity is a worldwide phenomenon and conservation of endemic species is becoming a pressing issue. However, species differ in life history characteristics, causing the best strategy for conservation to vary among species. Lucanus miwai is an endemic stag beetle of Taiwan, but the natural history and the conservation status of L. miwai have not been fully studied. Lucanus miwai adults live in forest-edge grassland habitats and are experiencing threats from anthropogenic exploitation. Additionally, climate change may threaten its long-term survival because of its specific habitat preference. Results from this study clearly indicate the need for more studies on the natural history of L. miwai and demonstrate that contemporary intensity of anthropogenic exploitation can have a negative effect on population size of this species. Future habitat loss because of global climate change, in addition, can be another major factor leading to population decline. Moreover, increasing demands on L. miwai because of population decline may lead to population extinction. In addition to protecting adult males from being collected, conservation strategies should also focus on discovering habitats utilized by individuals from different life stages, investigating the connectivity between populations, and maintaining the long-term availability of suitable habitats for L. miwai.     

Population field studies on the aphidophagous ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae): resource tracking and population characteristics

To clarify functional and numerical responses to temporal and spatial variations of resources (resource tracking), and the population characteristics of the ladybird beetle Harmonia axyridis, I analyzed the results of a 3-year field observation at 24 sites (seven plant species) on eight species of aphids. The seasonal changes in the number of beetles estimated by the Jolly–Seber method were significantly correlated with those of aphids in the total area. The estimated values of population parameters suggested frequent immigration and emigration of the H. axyridis population, although reproductive rates between spring and summer were rather stable all 3 years (1.87–3.49). The staying time and the daily number of adults and eggs at each site were influenced not only by a single factor but also by interactions among time and quantity and quality of the prey. The adult movement showed two patterns, which corresponded with the movement within and between the subpopulations when an assemblage of H. axyridis occurring on the plants of the same species or genus was regarded as a subpopulation. Adult movement intensely occurred within a subpopulation, although the beetles moving between subpopulations had a significantly greater chance to reach the habitat with a high aphid density. The habitats of H. axyridis could be categorized into a suitable habitat for survival and reproduction and a temporal refuge. The results obtained here suggest that H. axyridis, with high ability of prey searching and reproduction, maintains a stable population in heterogeneous and temporal habitats by its resource tracking mechanisms. Received: March 8, 1999 / Accepted: April 25, 2000     

Beetle and plant density as cues initiating dispersal in two species of adult predaceous diving beetles

Dispersal can influence population dynamics, species distributions, and community assembly, but few studies have attempted to determine the factors that affect dispersal of insects in natural populations. Consequently, little is known about how proximate factors affect the dispersal behavior of individuals or populations, or how an organism’s behavior may change in light of such factors. Adult predaceous diving beetles are active dispersers and are important predators in isolated aquatic habitats. We conducted interrelated studies to determine how several factors affected dispersal in two common pond-inhabiting species in southern Alberta, Canada: Graphoderus occidentalis and Rhantus sericans. Specifically, we (1) experimentally tested the effect of plant and beetle densities on dispersal probabilities in ponds; (2) surveyed ponds and determined the relationships among beetle densities and plant densities and water depth; and (3) conducted laboratory trials to determine how beetle behavior changed in response to variation in plant densities, conspecific densities, food, and water depth. Our field experiment determined that both species exhibited density dependence, with higher beetle densities leading to higher dispersal probabilities. Low plant density also appeared to increase beetle dispersal. Consistent with our experimental results, densities of R. sericans in ponds were significantly related to plant density and varied also with water depth; G. occidentalis densities did not vary with either factor. In the laboratory, behavior varied with plant density only for R. sericans, which swam at low density but were sedentary at high density. Both species responded to depth, with high beetle densities eliciting beetles to spend more time in deeper water. The presence of food caused opposite responses for G. occidentalis between experiments. Behavioral changes in response to patch-level heterogeneity likely influence dispersal in natural populations and are expected to be important for observed patterns of individuals in nature. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Distribution of ground-dwelling beetles in fragmented tropical habitats

The conservation of biodiversity is increasingly dependent on human-altered habitats. In a fragmented forest landscape in northern Costa Rica within the Mesoamerican Biological Corridor, an area of great conservation importance, we compared the diversity and composition of ground-dwelling beetle communities in five habitat types along a gradient of increasing disturbance: primary forest, logged forest, secondary forest, plantation (Gmelina arborea) and pasture. Using pitfall trapping we captured a total of 1,877 beetles (Coleoptera), comprising 422 morphospecies in 26 families. The plantation sites had the lowest number of species followed by secondary forest and pasture. Multivariate analysis separated the beetle fauna according to land use, and suggested that only the logged forest maintains a similar species assemblage to primary forest. However, each habitat harboured a number of unique species indicating the conservation value of the mosaic of habitats found in fragmented landscapes. Our results suggest that to maintain forest beetle species diversity it is essential that areas of natural forest are conserved. However, other land uses also provide beetle habitats and in fragmented areas active management to maintain a mosaic of land uses will contribute to the conservation of beetle diversity.     

Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape

The once extensive native forests of New Zealand’s central North Island are heavily fragmented, and the scattered remnants are now surrounded by a matrix of exotic pastoral grasslands and Pinus radiata plantation forests. The importance of these exotic habitats for native biodiversity is poorly understood. This study examines the utilisation of exotic plantation forests by native beetles in a heavily modified landscape. The diversity of selected beetle taxa was compared at multiple distances across edge gradients between each of the six possible combinations of adjacent pastoral, plantation, clearfell and native forest land-use types. Estimated species richness (Michaelis–Menten) was greater in production habitats than native forest; however this was largely due to the absence of exotic species in native forest. Beetle relative abundance was highest in clearfell-harvested areas, mainly due to colonisation by open-habitat, disturbance-adapted species. More importantly, though, of all the non-native habitats sampled, beetle species composition in mature P. radiata was most similar to native forest. Understanding the influence of key environmental factors and stand level management is important for enhancing biodiversity values within the landscape. Native habitat proximity was the most significant environmental correlate of beetle community composition, highlighting the importance of retaining native remnants within plantation landscapes. The proportion of exotic beetles was consistently low in mature plantation stands, however it increased in pasture sites at increasing distances from native forest. These results suggest that exotic plantation forests may provide important alternative habitat for native forest beetles in landscapes with a low proportion of native forest cover.     

Estimated density of four carabid beetle species in Mongolian oak and Korean red pine forest ecosystems on Baekdudaegan mountain range (Mt. Oade and Mt. Guryong) of South Korea

The estimation of animal population size is a primary field of interest for wildlife biologists, and such numerical estimation of wild animals is a very important factor in establishing national policy towards nature. Therefore, we performed this study to estimate the population density of carabid beetles preying on soil biota at the Mongolian oak (Quercus mongolica) forests and Korean red pine (Pinus densiflora) forests of two mountains, Mt. Odae and Mt. Guryong. We used the trapping web method with pitfall traps to collect the beetles. We calculated the estimated density of a total of four carabid beetle species in two specific regions and converted them to an estimated beetle population size within a certain area by using the study area. From our estimates, one beetle species, Leptocarabus seishinensis seishinensis L., displayed statistically significant results. Although there was no appreciable difference in the makeup of different carabid beetle species between the two forest communities, the population size of the beetle species was larger in Mongolian oak forests than in Korean red pine forests. The spatial distribution of carabid beetles by forest type did not show any preference for particular spots. Our results showed that carabid beetles did not have specific home ranges and that they were not density dependent. We expect to use the raw data to recognize the response of soil organisms to changes in the habitat environment and to track the patterns of change more accurately over long passages of time.     

Diversity, distinctiveness and conservation status of the Mediterranean coastal dung beetle assemblage in the Regional Natural Park of the Camargue (France)

The Mediterranean region as a whole has the highest dung beetle species richness within Europe. Natural coastal habitats in this region are among those which have suffered severe human disturbance. We studied dung beetle diversity and distinctiveness within one of the most important coastal protected areas in the west Euro‐Mediterranean region (the regional Park of Camargue, southern France) and made comparisons of dung beetle assemblages with other nearby Mediterranean localities, as well as with other coastal protected area (Doñana National Park, Spain). Our finding showed that: (1) The species richness of coastal habitats in the Camargue is low and only grasslands showed a similar level of species richness and abundance to inland habitats of other Mediterranean localities. The unique habitats of the coastal area (beaches, dunes and marshes) are largely colonized by species widely distributed in the hinterland. (2) In spite of their low general distinctiveness, dune and marsh edges are characterized by the occurrence of two rare, vulnerable, specialized and large roller dung beetle species of the genus Scarabaeus. As with other Mediterranean localities, current findings suggest a recent decline of Scarabaeus populations and the general loss of coastal dung beetle communities in Camargue. (3) The comparison of dung beetle assemblages between the Camargue and Doñana shows that, in spite of the low local dung beetle species richness in the Camargue, the regional dung beetle diversity is similar between both protected areas. Unique historical and geographical factors can explain the convergence in regional diversity as well as the striking divergence in the composition of dung beetle assemblages between both territories.     

A highly modified stygobiont diving beetle of the genus Copelatus (Coleoptera,Dytiscidae): taxonomy and cladistic analysis based on mitochondrial DNA sequences

Abstract. We report the discovery of a new species of Copelatus that is morphologically highly modified for life in groundwater. Copelatus abditus sp.n. was collected from a 4 m deep bore in central Australia. It was placed in Copelatinae based on morphological evidence. This is the first known stygobiont diving beetle that does not belong to the subfamily Hydroporinae. Sequences from the mitochondrial DNA cytochrome oxidase 1, 16S rRNA, tRNAL and NADH dehydrogenase subunit 1 genes suggest that the species belongs to the subgenus Papuadytes, the morphological apomorphies of which are reduced in the new groundwater species. Copelatus (Papuadytes) abditus very much resembles other world stygobiont dytiscids, which all belong to the subfamily Hydroporinae. We suggest that this striking convergence is due to similar selective pressures imposed on all groundwater beetles. We suggest a scenario in which species from temporary habitats evade drought by entering the hyporheic zone, and may be driven to a subterranean existence by desertification of their ancestral habitats, as can be observed in Australia.     

Parasitism and olfactory responses of <Emphasis Type="Italic">Dastarcus helophoroides</Emphasis> (Coleoptera: Bothrideridae) to different Cerambycid hosts

Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae) is an important natural enemy of longhorned beetles (Coleoptera: Cerambycidae). It is distributed throughout most Provinces in China. We investigated whether there were differences among D. helophoroides populations collected from different hosts in different geographic locations. Results showed that different D. helophoroides populations displayed different olfactory responses to larval frass from different longhorned beetle species. All populations were significantly attracted to the frass of their original hosts. Parasitism rates of different populations also varied when supplied with host larvae of the same longhorned beetle species. These results indicate that the three D. helophoroides populations tested differed in host-related behaviors. Therefore, the population of D. helophoroides must be taken into consideration when implementing biological control programs for different species of longhorned beetle.