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.

     

Colonization of disturbed Scots pine trees by bark‐ and wood‐boring beetles

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Synchronous rise and fall of bark beetle and parasitoid populations in windthrow areas

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Behaviours of phoretic mites (Acari) associated with Ips pini and Ips grandicollis (Coleoptera: Curculionidae) during host‐tree colonization

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Ethanol‐injection induces attacks by ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) on a variety of tree species

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Role of two phoretic mites in transmission of bluestain fungus, Ceratocystis minor

ABSTRACT.

• 1 Tarsonemus mites phoretic on southern pine beetles were shown to be hypervectors of bluestain fungus, Ceratocystis minor (Hedgcock 1906).
• 2 Beetles with phoretic mites had significantly more C.minor than did mite-free beetles.
• 3 Over 90% of tarsonemid mites collected from beetle-infested inner bark and 59% of mites phoretic on emerging beetles carried C.minor spores.
• 4 This association is an interesting example of a three-way mutualism among insects, mites and fungi.

     

Evidence of interaction between Sirex noctilio and other species inhabiting the bole of Pinus

• 1 Sirex noctilio is a woodwasp native to Eurasia and Northern Africa and has recently been found infesting pines in eastern North America. Its pest status in this new range is not yet known, although it is an important pest in other areas where it has been introduced. Pinus spp. in North America are hosts to several native and alien species of subcortical insects. Interactions between the woodwasp and these species may influence its distribution or the characteristics of its life history, thus affecting its population dynamics over time.
• 2 Sixty S. noctilio‐infested Pinus spp. were felled in Ontario, Canada, and all phloem‐feeding and woodboring insects were collected and identified from each 1‐m section of the tree.
• 3 Sirex noctilio was in a tree alone 10% of the time but commonly shared the tree with subcortical beetles, such as Tomicus piniperda, Pissodes nemorensis, Ips grandicollis, Gnathotrichus materiarius and Monochamus carolinensis. The woodwasp was distributed throughout the tree stem and this distribution overlapped with that of the beetles.
• 4 Fewer but larger S. noctilio males emerged from trees with beetles compared with those without, although there was no statistical difference in females.
• 5 These findings suggest that co‐habiting beetles could negatively affect S. noctilio population dynamics over time. The potential mechanisms for this interaction are discussed.

     

Association of Geosmithia fungi (Ascomycota: Hypocreales) with pine- and spruce-infesting bark beetles in Poland

Bark beetles (Coleoptera, Scolytinae) are known to be associated with fungi, particularly species of the orders Ophiostomatales and Microascales. However, very little is known about other ectosymbionts of phloeophagous bark beetles on Pinaceae. In this study, we examined the Geosmithia species associated with eight bark beetle species infesting Picea abies and Pinus sylvestris branches in Poland. Fungi were isolated from 1 731 samples collected from 14 study sites. We identified a total of 653 isolates that were sorted into nine taxa based on their phenotypic similarity and phylogeny of their ITS-LSU regions of rDNA, β-tubulin, elongation factor 1α and the second-largest subunit of the RNA polymerase II gene. They represented nine species without formal names. There were large quantitative and qualitative differences in the composition of Geosmithia communities between P. sylvestris and P. abies trees. The proportion of samples infested with Geosmithia species suggests that this association is more widespread among bark beetles infesting branches of P. sylvestris. In addition, these beetles were vectors of different Geosmithia species compared with than the beetles that colonize P. abies. In mixed-conifer forests, the Geosmithia communities were more diverse and richer than in pure spruce or pine stands, where the insects Pityogenes chalcographus and Pityophthorus pityographus with low host-specificity play a distributing role for various Geosmithia species. Among eight bark beetle species examined, only P. bidentatus, P. pityographus, P. chalcographus and Polygraphus poligraphus acted as effective vectors for Geosmithia species. The following hypothesis emerges from these studies: changes in the composition of ectosymbionts of pine- and spruce-infesting bark beetles in Central Europe run along a gradient of thickness of the wood substrata preferred by insects.     

Phoretic mites and nematode associates of Scolytus multistriatus and Scolytus pygmaeus (Coleoptera: Scolytidae) in Austria

Abstract 1 The species assemblages and abundance of phoretic mites and nematodes associated with the elm bark beetles, Scolytus multistriatus and Scolytus pygmaeus, were studied in Austria. 2 A total of 3922 individual mites were recorded from 144 adults of S. multistriatus and 178 adults of S. pygmaeus. The species spectrum was identical and the relative abundance of mites was very similar for both species of scolytids. Nine mite species, Pyemotes scolyti, Pseudotarsonemoides eccoptogasteri, Trichouropoda bipilis, Tarsonemus crassus, Proctolaelaps eccoptogasteris, Proctolaelaps scolyti, Chelacheles michalskii, nr. Eueremaeus sp. and Elattoma sp. were detected. Two of the nine species, nr. Eueremaeus sp. and Elattoma sp., are documented here as new associates of Scolytus spp. 3 Pyemotes scolyti was the most frequent mite species, and Ps. eccoptogasteri and T. bipilis were relatively common, whereas the other mites occurred occasionally or were rare. 4 The trophic roles of most of the mites associated with S. multistriatus and S. pygmaeus are poorly known, but they may include fungivores, parasitoids of bark beetle broods, predators of bark beetle broods and/or mites and/or nematodes. 5 Besides phoretic mites, two nematode associates were seen on the investigated insects. A species of Cryptaphelenchus occurred under the elytra of both scolytid species, whereas the adults of a Neoparasitylenchus sp. were present inside abdomens of S. multistriatus, but absent from S. pygmaeus.     

The first fossil record of mites of the family Pyemotidae (Acari: Heterostigmata), with description of a new species of the genus <Emphasis Type="Italic">Pyemotes</Emphasis> from Rovno amber

The first fossil mite of the family Pyemotidae (Acari: Heterostigmata) is recorded. Pyemotes primus sp. nov. is described from the Late Eocene of the Rovno amber from a syninclusion with the bark beetles Taphramites rovnoensis Petrov et Perkovsky, 2008. In the presence of well developed longitudinal striation on hysterosomal tergites, the new species is similar to species from scolyti group, which is characterized by parasitoidism on immature stages of bark beetles and phoresy on adult stages. P. primus differs from the closely related species P. dryas (Vitzthum, 1923), P. parviscolyti Cross et Moser, 1971, P. johnmoseri Khaustov, 2004, and P. mandelshtami Khaustov, 1998 in the subequal setae h ₁ and h ₂.     

Defence syndromes in lodgepole – whitebark pine ecosystems relate to degree of historical exposure to mountain pine beetles

Warming climate is allowing tree‐killing bark beetles to expand their ranges and access naïve and semi‐naïve conifers. Conifers respond to attack using complex mixtures of chemical defences that can impede beetle success, but beetles exploit some compounds for host location and communication. Outcomes of changing relationships will depend on concentrations and compositions of multiple host compounds, which are largely unknown. We analysed constitutive and induced chemistries of Dendroctonus ponderosae's primary historical host, Pinus contorta, and Pinus albicaulis, a high‐elevation species whose encounters with this beetle are transitioning from intermittent to continuous. We quantified multiple classes of terpenes, phenolics, carbohydrates and minerals. Pinus contorta had higher constitutive allocation to, and generally stronger inducibility of, compounds that resist these beetle–fungal complexes. Pinus albicaulis contained higher proportions of specific monoterpenes that enhance pheromone communication, and lower induction of pheromone inhibitors. Induced P. contorta increased insecticidal and fungicidal compounds simultaneously, whereas P. albicaulis responses against these agents were inverse. Induced terpene accumulation was accompanied by decreased non‐structural carbohydrates, primarily sugars, in P. contorta, but not P. albicaulis, which contained primarily starches. These results show some host species with continuous exposure to bark beetles have more thoroughly integrated defence syndromes than less‐continuously exposed host species.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

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.     

Comparisons of mountain pine beetle (Dendroctonus ponderosae Hopkins) reproduction within a novel and traditional host: effects of insect natal history,colonized host species and competitors

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Variables associated with the occurrence of Ips beetles,red turpentine beetle and wood borers in live and dead ponderosa pines with post‐fire injury

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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.     

New associations of phoretic mites on <Emphasis Type="Italic">Pityokteines curvidens</Emphasis> (Coleoptera,Curculionidae, Scolytinae)

The species composition and abundance of phoretic mites of the bark beetle Pityokteines curvidens caught in pheromone traps were investigated in Croatia. The P. curvidens trapping programs have been in an experimental phase in Croatia since 2004 as a possible monitoring and control system. The trapping program also permits the opportunity to sample phoretic mites found associated with the beetles. Beetles were caught using Curviwit pheromones in Theysohn traps placed in the Litorić region of Croatia. A total of 12 mite species were recovered, including Schizostethus simulatrix, Dendrolaelaps quadrisetus, Histiostoma piceae, H. cf. varia, Paraleius leontonychus, Pleuronectocelaeno barbara, Tarsonemus minimax, Trichouropoda lamellosa, Uroobovella ipidis, Schwiebea sp., Phauloppia lucorum and Dolicheremaeus dorni. Five species, Pl. barbara, Schwiebea sp., H. cf. varia, Ph. lucorum and Do. dorni, are identified for the first time in association with P. curvidens. These findings increase the number of mite species known to be phoretic on P. curvidens from 11 to 16. The present study also increases the number of known mite associates of Pityokteines spp. from 14 to 18.     

Interaction of pre-attack and induced monoterpene concentrations in host conifer defense against bark beetle-fungal complexes

Two pine species (Pinus resinosa, P. banksiana) responded to inoculation with fungi carried by bark beetles by rapidly increasing monoterpene concentrations at the entry site. Changes in total monoterpenes were more pronounced than changes in proportionate compositions. The extent and rate of host response was affected by fungal species, the viability of the inoculum, and host tree species. In general, host responses were highest to fungi that are phytopathogenic and consistently associated with the major bark beetles in the study region. Simple mechanical wounding cannot account for the observed allelochemical changes, as aseptic inoculations elicited only minor reactions. Similarly, inoculation with autoclaved inviable fungi generally elicited intermediate responses, suggesting that both structural and metabolic fungal properties are important. Responses by jack pine, P. banksiana, were generally more rapid and variable than those of red pine, P. resinosa. Dose-toxicity experiments with synthetic compounds demonstrated that monoterpene concentrations present in vivo only a few days after simulated attack are lethal to most beetles. Constitutive (pre-attack) monoterpene levels can also exert some toxicity. Because bark beetles engage in pheromone-mediated mass attacks that can deplete host defenses, constitutive monoterpene levels, while a necessary early phase of successful plant defense, appear insufficient by themselves. Such interactions between constitutive and induced defense chemistry may be important considerations when evaluating general theories of plant defense.     

Phoretic Poecilochirus mites specialize on their burying beetle hosts

Recurring species interactions can cause species to adapt to each other. Specialization will increase the fitness of symbionts in the coevolved association but may reduce the flexibility of symbiont choice as it will often decrease fitness in interactions with other than the main symbiont species. We analyzed the fitness interactions between a complex of two cryptic mite species and their sympatric burying beetle hosts in a European population. Poecilochirus mites (Mesostigmata, Parasitidae) are phoretic on burying beetles and reproduce alongside beetles, while these care for their offspring at vertebrate carcasses. While Poecilochirus carabi is typically found on Nicrophorus vespilloides beetles, P. necrophori is associated with N. vespillo. It has long been known that the mites discriminate between the two beetle species, but the fitness consequences of this choice remained unknown. We experimentally associated both mite species with both beetle species and found that mite fitness suffered when mites reproduced alongside a nonpreferred host. In turn, there is evidence that one of the beetle species is better able to cope with the mite species they are typically associated with. The overall fitness effect of mites on beetles was negative in our laboratory experiments. The Poecilochirus mites studied here are thus specialized competitors or parasites of burying beetles.