Behavior of male and female spruce bark beetles,Ips typographus,on the bark of host trees during mass attack

The behavior of 118 spruce bark beetles, Ips typographus,was observed on trees under colonization. Most individuals were followed from when they landed until they entered or left the tree. Both males and females spent most time inspecting crevices and searching for a place to start boring or for a hole to enter. These behaviors accounted for 87 and 70% of all behavioral acts recorded for males and females, respectively. Females entered galleries with males only after a period of pushing at the gallery entrance. Males spent on average 3 min and females 4 min on the bark before entering or leaving the tree. Thirty-three percent of the beetles eventually entered the tree, 31% flew away, 35% dropped from the host, and one beetle was eaten by a predator. The results are discussed in relation to the question of mate choice in bark beetles and to studies on attack dynamics of spruce bark beetle populations.     

Ophiostomatoid fungi isolated from Japanese red pine and their relationships with bark beetles

We isolated ophiostomatoid fungi from bark beetles infesting Pinus densiflora and their galleries at 24 sites in Japan. Twenty-one ophiostomatoid fungi, including species of Ophiostoma, Grosmannia, Ceratocystiopsis, Leptographium, and Pesotum, were identified. Among these, 11 species were either newly recorded in Japan or were previously undescribed species. Some of these fungal species were isolated from several bark beetles, but other species were isolated from only a particular beetle species. Thus, it is suggested that some ophiostomatoid fungi have specific relationships with particular beetle species. In addition, fungus-beetle biplots from redundancy analysis (RDA) summarizing the effects of beetle ecological characteristics suggested that the association patterns between bark beetles and the associated fungi seemed to be related to the niches occupied by the beetles.     

Pine monoterpenes and pine bark beetles: a marriage of convenience for defense and chemical communication

Pine-feeding bark beetles (Coleoptera: Scolytidae) interact chemically with their host pines (Coniferales: Pinaceae) via the behavioral, physiological, and biochemical effects of one class of isoprenoids, the monoterpenes and their derivatives. Pine monoterpenes occur in the oleoresin and function as behaviorally active kairomones for pine bark beetles and their predators, presenting a classic example of tri-trophic chemical communication. The monoterpenes are also essential co-attractants for pine bark beetle aggregation pheromones. Ironically, pine monoterpenes are also toxic physiologically to bark beetles at high vapor concentrations and are considered an important component of the defense of pines. Research over the last 30 years has demonstrated that some bark beetle aggregation pheromones arise through oxygenation of monoterpenes, linking pheromone biosynthesis to the host pines. Over the last 10 years, however, several frequently occurring oxygenated monoterpene pheromone components (e.g., ipsenol, ipsdienol and frontalin) have also been shown to arise through highly regulated de novo pathways in the beetles (reviewed in Seybold and Tittiger, 2003). The most interesting nexus between these insects and their plant hosts involves the late-stage reactions in the monoterpenoid biosynthetic pathway, during which isomeric dimethylallyl diphosphate and isopentenyl diphosphate are ultimately elaborated to stereospecific monoterpenes in the trees and to hydroxylated monoterpenes or bicyclic acetals in the insects. There is signal stereospecificity in both production of and response to the monoterpenoid aggregation pheromones of bark beetles and in response to␣the monoterpenes of the pines. In the California fivespined ips, Ips paraconfusus, we have discovered a number of cytochome P450 genes that have expression patterns indicating that they may be involved in detoxifying monoterpene secondary metabolites and/or biosynthesizing pheromone components. Both processes result in the production of oxygenated monoterpenes, likely with varying degrees of stereospecificity. A behavioral analysis of the stereospecific response of I. paraconfusus to its pheromone is providing new insights into the development of an efficacious bait for the detection of this polyphagous insect in areas outside the western United States. In contrast, a Eurasian species that has arrived in California, the Mediterranean pine engraver, Orthotomicus (Ips) erosus, utilizes both a monoterpenoid (ipsdienol) and a hemiterpenoid (2-methyl-3-buten-2-ol) in its pheromone blend. The stereospecificity of the response of O. erosus to the monoterpenoid appears to be the key factor to the improved potency of the attractant bait for this invasive species.Dedicated to Professor David L. Wood on the occasion of his 75th birthday, January 8, 2006     

Helicosporidium infection of the great European spruce bark beetle, Dendroctonus micans (Coleoptera: Scolytidae)

An infection of the great European spruce bark beetle Dendroctonus micans (from Turkey) by the parasitic green alga Helicosporidium is described. This is the first time that Helicosporidium has been found to infect a bark beetle (Scolytidae) and the first time that a naturally infected beetle has been reported for the Eurasian continent. The typical cysts of Helicosporidium contain three ovoid cells and one helical, filamentous cell. The morphological characteristics are revealed by light and electron microscopy. Distinct electron-dense inclusions with a peculiar ultrastructure may represent pyrenoids. Since D. micans is an important pest, the discovery of a natural pathogen may offer a chance for biological control.     

Positional behavior ofCebuella pygmaea in Yasuni National Park, Ecuador

In terms of positional behavior, the small-bodied callitrichids are distinguished from other anthropoids by their ability to leap between and to cling to large vertical supports, feeding occasionally or frequently on sap and bark insects. In this paper, I studied the positional behavior of a group of pygmy marmosets in a hilly wet tropical forest in Yasuni National Park, Ecuador. During traveling, the animals used quadrupedal walk/bound and leaping mostly on small horizontal supports. During foraging, quadrupedal walk/bound and clambering dominated. The main foraging postures were stand and cantilever, occurring mainly on small horizontal lianas. During feeding, scansorial locomotion was used very frequently. Claw clinging was the dominant feeding posture. Furthermore, large vertical lianas and tree boles were the most frequent feeding supports. These observations would suggest that scansorial locomotion, vertical clinging, and vertical leaping are most likelynot part of the same form function complex. Vertical leaps appear to be associated with moving in the lower parts of the forest. On the other hand, scansorial locomotion and vertical clinging appear to be related to feeding on the sap of tree boles and large lianas in the lower parts of the forest.     

Extreme ecological stoichiometry of a bark beetle–fungus mutualism

1. Ecological stoichiometry theory was applied to investigate how a consumer contends with an extreme elemental mismatch between its food and its body via symbiotic facilitation. 2. The beetle Dendroctonus brevicomis LeConte develops in bark, a substrate extremely low in nitrogen (N) and phosphorus (P). Its survival there depends on interactions with mutualist and antagonist fungi. 3. This study found that mutualists transfer N and P from sapwood and phloem into bark, where beetles feed, whereas the antagonist moves these elements only to phloem, resulting in starvation of the insect. However, even with mutualists, N and P concentrations remained low in bark, resulting in low N and extremely low P concentrations in the beetle. 4. The N:P ratios found in D. brevicomis larvae were the highest thus far reported for beetles and among the highest for insects and invertebrates. This suggests that the beetle has evolved additional, nutrient‐sparing adaptations.     

<Emphasis Type="Italic">Geosmithia</Emphasis> Fungi are Highly Diverse and Consistent Bark Beetle Associates: Evidence from their Community Structure in Temperate Europe

Geosmithia spp. (Ascomycota: Hypocreales) are little-studied, dry-spored fungi that occur in galleries built by many phloeophagous bark beetles. This study mapped the distribution and environmental preferences of Geosmithia species occurring in galleries of temperate European bark beetles. One hundred seven host tree samples of 16 tree species infested with 23 subcortical insect species were collected from across Europe during the years 1997–2005. Over 600 Geosmithia isolates from the beetles were sorted into 17 operational taxonomic units (OTUs) based on their phenotype similarity and phylogeny of internal transcribed spacer (ITS) region of rDNA (ITS1-5.8S-ITS2). The OTUs represent six known species and eight undescribed taxa. Ninety-two samples infested with subcortical insects were characterized by the presence/absence of OTUs and the similarity among the samples was evaluated. Geographically distant populations of the same beetle species host relatively uniform Geosmithia communities across large geographic areas (ranging from southern Bulgaria to the Czech Republic). This suggests effective dispersal of Geosmithia spp. by bark beetles. Clustering of similar samples in ordination analysis is correlated predominantly with the isolation source (bark beetles and their respective feeding plant), but not with their geographical origin. The composition of the Geosmithia OTU community of each bark beetle species depends on the degree of isolation of the species’ niches. Thus, Geosmithia communities associated with regularly co-occurring bark beetle species are highly similar. The similarity decreases with decreasing frequency of beetle species’ co-occurrence, a pattern resembling that of entomochoric ophiostomatoid fungi. These findings suggest that: 1) communities of Geosmithia spp. are vector-specific; 2) at least in some cases, the association between Geosmithia OTUs and bark beetles may have been very stable and symbioses are likely to be a fundamental factor in the speciation of Geosmithia fungi; and 3) that even nonsticky spores of Geosmithia are suitable for maintaining an insect–fungus association, contrary to previous hypotheses. An erratum to this article can be found at     

Differences in the fruit maturation stages at which oviposition occurs among insect seed predators feeding on the fruits of five dipterocarp tree species

The seeds of dipterocarp trees are the main food resources for many species of weevils, bark beetles and small moths; however, for most seed‐eating insects on dipterocarp tropical trees, seed utilization patterns remain poorly investigated. This study aimed to determine the fruit maturation stages at which eggs are laid by different insect seed predators feeding on the seeds or fruits of the following five dipterocarp species: Dipterocarpus globosus, Dryobalanops aromatica, Shorea beccariana, S. acuta and S. curtisii, which reproduced during the same period. We investigated the occurrence frequencies of the insect seed predators at various growth stages by collecting both unfallen and fallen fruit on several occasions during the period of seed/fruit maturation in a tropical rainforest in Borneo from September to December 2013. Weevils and bark beetles were the dominant insect seed predators of the five tree species. One or two weevil species of Alcidodes, Damnux and/or Nanophyes preyed on the seeds of each of the five tree species, and one bark beetle species, Coccotrypes gedeanus, preyed on the seeds of all five tree species. Many larvae, pupae and adults of each weevil species were found in pre‐dispersal (unfallen) fruit, whereas bark beetles at various growth stages were found in post‐dispersal (fallen) fruit. These results suggested that, among the dominant insect seed predators of the five dipterocarp species, weevil species oviposit on pre‐dispersal fruit and begin their larval growth before seed dispersal, whereas the oviposition and larval development of bark beetle species occurs in post‐dispersal fruit.     

Conifer phenolic resistance markers are bark beetle antifeedant semiochemicals

1 Phenols are important in conifer resistance to fungi associated with bark beetles and as markers for resistance to beetle mass‐attacks. However, the mechanism of phenolic activity in conifer resistance to Ips typographus bark beetle remains unclear. 2 By a new bioassay, we tested the effect on host acceptance and tunnelling of male and female I. typographus in an artificial gallery (13 mm long) for 4 h (no‐choice test). To the artificial diet, an aliquot of host (catechin, taxifolin, or resveratrol) or nonhost (E‐conophthorin) compounds was added, singly tested at doses close to those of tree bark. Host acceptance and tunnelling was measured by the amount of diet removed by the insects. 3 All tested chemicals decreased the tunnelling activity of I. typographus, with an antifeedant effect stronger in males and increasing with dose. No mortality was recorded. The nonhost volatile spiroketal, E‐conophthorin, had the highest antifeedant activity both in males and females. Among host compounds, effects and dose–response were weak in females. Both catechin and E‐conophthorin gave a 50% reduction of tunnelling at a concentration of 0.1% for males, the pioneering (host selecting) sex in Ips. The threshold of activity for host compounds to males was at concentrations of 0.03–0.1%, which corresponds to, or is less than, the concentrations reported from spruce host bark. 4 The results allow us to support the hypothesis of a direct behavioural antifeedant mechanism for resistance from those phenols that are particularly active for the pioneering males during tree attack.     

Mite communities (Acari,Mesostigmata) associated with <Emphasis Type="Italic">Ips typographus</Emphasis> (Coleoptera,Scolytidae) in managed and natural Norway spruce stands in Central Europe

Previous studies have suggested that forest management practices can influence bark beetle populations as well as those of some associated insects. However, the impact on bark beetle-associated mites, which occur in bark beetle galleries in large numbers, have not yet been studied. The objective of this study was to compare mesostigmatid mite communities associated with the Norway spruce pest Ips typographus in managed and natural forest stands separated by spruce-free belt. The study sites were located in Białowieża National Park (NE Poland) as well as in the Izery Mountains (Szklarska Poręba Forest District — SW Poland), which were destroyed in 1981–1987 by an ecological disaster. In total, 30 Borregard pipe traps containing the commercial attractant Ipsodor W (Chemipan, Poland) were set up at each study site and collected in August 2010. In total, 7214 bark beetles and 1804 mites were collected which were classified into 16 species. We observed differences in the total abundance of mites as well as the total number of recorded mite species. The communities were quite similar, and were generally dominated by populations of Trichouropoda polytricha, Dendrolaelaps quadrisetus and Uroobovella ipidis. The Shannon and Evenness indexes as well as the mean number of mites per sample were not significantly different between forests.     

Measurement of the sieve tube membrane potential

A procedure is described for the measurement of the sieve tube membrane potential in the phloem of bark strips from Salix exigua Nutt. Measurements were made by inserting a measuring microelectrode into sap exuding from severed stylets of the willow aphid, Tuberolachnus salignus. Data taken from 20 bark strips gave an average potential of −155 ± 9 millivolts. Evidence is presented for an electrogenic component of the sieve tube membrane potential. The occurrence of a saturable sucrose-induced membrane depolarization is consistent with the concept of sugar accumulation by a sucrose/H⁺ co-transport mechanism.     

Bark beetle and woodborer responses to stand thinning and prescribed fire in northeastern US coastal and inland pitch pine barrens

1. Southern pine beetle, Dendroctonus frontalis, has expanded its range further into the northeastern United States. This expansion threatens rare and ecologically valuable interior and coastal pitch pine barrens.
2. Pitch pine barrens restoration and southern pine beetle infestation suppression often involve leaving downed dead wood that saproxylic insects can exploit.
3. Semiochemical-baited traps were used to investigate the response of bark beetles and woodborers to restoration treatments at Rocky Point State Forest and the Albany Pine Bush Preserve, examples of coastal and interior pitch pine forests, respectively.
4. A total of 29,598 saproxylic insects from 116 species of bark beetles and woodborers were captured at Rocky Point State Forest, while 23,117 individuals from 67 species were captured at Albany Pine Bush Preserve.
5. Ips spp. were abundant at both sites with 28%–47% and 42%–74% of total collections at Rocky Point State Forest and Albany Pine Bush Preserve, respectively.
6. Ips grandicollis did not respond to treatments at either site. However, Ips pini was found in higher numbers in thinned blocks in Rocky Point State Forest.

     

Interspecific relationship between the Japanese horned beetle and two Japanese stag beetle species

The Japanese horned beetle Trypoxylus dichotomus septentrionalis and the Japanese stag beetles Lucanus maculifemoratus maculifemoratus and Prosopocoilus inclinatus inclinatus generally occur syntopically and aggregate on oak tree surfaces that exude sap. Securement of these sap sites might be directly linked to male reproductive success. Among the three species, it is likely that males of T. d. septentrionalis are dominant in occupation of feeding sites because of their larger body size. However, there is no clear evidence of this superiority. Moreover, if T. d. septentrionalis is dominant, the mechanism by which the two stag beetle species secure the feeding sites remains unclear. In the present study, I used body mass to compare the body size among males of T. d. septentrionalis, L. m. maculifemoratus and P. i. inclinatus. Further, to clarify the interspecific relationship between the horned beetle and the two stag beetles, I investigated the seasonality of emergence pattern of males at the feeding sites in the field. Comparison of body mass and observation of fighting behavior revealed that males of T. d. septentrionalis have an apparent superiority over males of the two stag beetle species. The seasonal emergence patterns of the two stag beetle species at the feeding sites showed bimodal distributions, and avoided the peak of emergence in T. d. septentrionalis. My results suggest that the two stag beetle species exhibit mate‐securing tactics by emerging at the feeding sites early and late during the breeding season, in order to avoid encountering T. d. septentrionalis.     

Does Rust Infection of Willow Affect Feeding and Oviposition Behavior of Willow Leaf Beetles?

Willows are often attacked by both herbivorous insects and rust fungi. Little is known about interactions between these two willow enemies. We studied whether feeding and oviposition behavior of the willow leaf beetle Plagiodera versicolora upon the willow hybrid Salix x cuspidata is affected when the rust fungus Melampsora allii-fragilis has attacked the plant. Laboratory bioassays revealed that adult willow leaf beetles significantly avoided feeding and oviposition on rust-infected leaves when compared to healthy leaves. Further bioassays aimed to elucidate the temporal and spatial scale of effects of rust infection on feeding behavior of adults. While infected parts of leaves were avoided at all times past infection tested (8, 12, and 16 days), symptom-free parts of infected leaves were only avoided 16 days past infection. Systemic effects extended only one leaf position up and two leaf positions down from the infection site.     

Bark beetles as lidar targets and prospects of photonic surveillance

Forestry is raising concern about the outbreaks of European spruce bark beetle, Ips typographus, causing extensive damage to the spruce forest and timber values. Precise monitoring of these beetles is a necessary step towards preventing outbreaks. Current commercial monitoring methods are catch-based and lack in both temporal and spatial resolution. In this work, light scattering from beetles is characterized, and the feasibility of entomological lidar as a tool for long-term monitoring of bark beetles is explored. Laboratory optical properties, wing thickness, and wingbeat frequency of bark beetles are reported, and these parameters can infer target identity in lidar data. Lidar results from a Swedish forest with controlled bark beetle release event are presented. The capability of lidar to simultaneously monitor both insects and a pheromone plume mixed with chemical smoke governing the dispersal of many insects is demonstrated. In conclusion, entomological lidar is a promising tool for monitoring bark beetles.     

Confinement Behavior of Cape Honey Bees (Apis mellifera capensis Esch.) in Relation to Population Densities of Small Hive Beetles (Aethina tumida Murray)

We quantified the effects of increasing small hive beetle (Aethina tumida Murray) populations on guarding behavior of Cape honey bees (Apis mellifera capensis, an African subspecies). We found more confinement sites (prisons) at the higher (50 beetles per colony) rather than lower (25 beetles per colony) beetle density. The number of beetles per prison did not change with beetle density. There were more guard bees per beetle during evening than morning. Neither guard bee nor beetle behavior varied with beetle density or over time. Forty-six percent of all beetles were found among the combs at the low beetle density and this increased to 58% at the higher one. In neither instance were beetles causing depredation to host colonies. Within the limits of the experiment, guarding behavior of Cape honey bees is relatively unaffected by increasing beetle density (even if significant proportions of beetles reach the combs).     

Tracking down the "head blob": comparative analysis of wingless expression in the developing insect procephalon reveals progressive reduction of embryonic visual system patterning in higher insects

The evolution of larval head morphology in holometabolous insects is characterized by reduction of antennal appendages and the visual system components. Little insight has been gained into molecular developmental changes underlying this morphological diversification. Here we compare the expression of the segment polarity gene wingless (wg) in the pregnathal head of fruit fly, flour beetle and grasshopper embryos. We provide evidence that wg activity contributes to segment border formation, and, subsequently, the separation of the visual system and protocerebrum anlagen in the anterior procephalon. In directly developing insects like grasshopper, seven expression domains are formed during this process. The activation of four of these, which correspond to polar expression pairs in the optic lobe anlagen and the protocerebral ectoderm, has shifted to postembryonic stages in flour beetle and Drosophila. The remaining three domains map to the protocerebral neuroectoderm, and form by disintegration of a large precursor domain in flour beetle and grasshopper. In Drosophila, the precursor domain remains intact, constituting the previously described “head blob”. These data document major changes in the expression of an early patterning gene correlated with the dramatic evolution of embryonic visual system development in the Holometabola.     

Anomalous outbreaks of an invasive defoliator and native bark beetle facilitated by warm temperatures,changes in precipitation and interspecific interactions

Biotic disturbance agents such as insects can be highly responsive to climatic change and have widespread ecological and economic impacts on forests. Quantifying the responses of introduced and native insects to climate, including how dynamics of one agent may mediate those of another, is important for forecasting disturbance and associated impacts on forest structure and function. We investigated drivers of outbreaks by larch casebearer Coleophora laricella, an invasive defoliator, and eastern larch beetle Dendroctonus simplex, a native, tree‐killing bark beetle, on tamarack Larix laricina from 2000 to in Minnesota, USA. We evaluated the utility of temporal, spatial and climatic variables in predicting the presence/absence of outbreaks of each insect in cells of rasterized aerial survey data. The role of defoliation by larch casebearer in outbreaks of eastern larch beetle was also investigated. For both species, the most important predictors of outbreak occurrence were proximity of conspecific outbreaks in space and time. For larch casebearer, outbreak occurrence was positively associated with spring precipitation and warmer growing seasons. Outbreak occurrence of eastern larch beetle was positively associated with warmer and dryer years and was more likely in cells with prior defoliation by larch casebearer. Our results demonstrate that climate can drive large scale outbreaks of introduced and non‐native disturbance agents on a single host species, and that interactions at the tree level between such agents may scale up to manifest across large temporal and spatial scales.     

Kairomonal Responses of Natural Enemies and Associates of the Southern Ips (Coleoptera: Curculionidae: Scolytinae) to Ipsdienol, Ipsenol and Cis-Verbenol

Bark beetle infested pines are an ephemeral habitat utilized by a diverse assemblage of insects. Although many bark beetle insect associates have little or no measurable impact on bark beetle brood production, some reduce brood production by either competing with brood for the limited phloem tissue or by feeding on brood. Several studies have observed synchrony between the colonization of hosts by bark beetles and the arrival of insect associates. Some insect associates mediate synchrony with bark beetle mass attacks with kairomonal responses to bark beetle aggregation pheromones. The objectives of this study were to document the community of Coleoptera associated with the southern Ips (Ips avulsus, Ips calligraphus and Ips grandicollis) and to test the hypothesis that synchrony of insect associates with the southern Ips is mediated by kairomonal responses to aggregation pheromones. A large community of Coleoptera (109 species) was recorded from traps baited with southern Ips pheromones. A significant treatment effect was observed for the guilds of meristem feeders, natural enemies and woodborers. The southern Ips pheromone ipsenol was broadly attractive to meristem feeders, natural enemies and woodborers and in general blends were more attractive than individual compounds. These results demonstrate that a diverse community of Coleoptera is associated with the southern Ips and that several members of this community facilitate synchrony with kairomonal responses to southern Ips aggregation pheromones.     

Resource Partitioning among Parasitoids (Hymenoptera: Braconidae) of Phoracantha semipunctata in Their Native Range

The Eucalyptus longhorned borer, Phoracantha semipunctata (F.), is native to Australia, but it has been introduced without its natural enemies into many parts of the world in which its Eucalyptus spp. host has been planted. The beetle has developed large populations in these novel habitats and has been responsible for the mortality of large numbers of trees. Although there is a considerable catalogue of the parasitoids of the beetle in Australia, limited ecological information on the assemblage of parasitoids attacking P. semipunctata is available. We removed bark from 40 felled trees, recorded gallery width and bark thickness over parasitized larvae, and removed all parasitoids. Adult size, sex, and species were recorded when the parasitoid pupae eclosed. Syngaster lepidus Brullè, Jarra phoracantha Austin, Quicke, and Marsh, J. maculipennis Austin, Quicke, and Marsh, and J. painei Austin and Dangerfield were most commonly collected. The solitary parasitoid S. lepidus preferred smaller larvae than did the gregarious Jarra spp. The two species with shorter ovipositors, J. maculipennis and J. painei, parasitized larvae under thinner bark than did the other two species with longer ovipositors. There was a significant positive correlation between host larval size and number of parasitoid pupae of the gregarious species. Also, there was a significant positive correlation between host larval size and parasitoid adult size. The ecological relationships between this assemblage of parasitoids and their beetle host may be useful in establishing an effective biological control program.