Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Temperature-dependent reproduction of the spruce bark beetle Ips typographus,and analysis of the potential population growth

1. The spruce bark beetle Ips typographus (L.) is one of the most important forest pests in Central Europe, but despite this the effects of temperature on life history and population growth are largely unknown. This study examines the effects of temperature on reproduction and intrinsic demographic statistics. 2. Laboratory experiments on oviposition were carried out at six temperatures in the range 12–33 °C, using the so-called sandwich rearing technique for bark beetles. 3. A linear relationship between oviposition rates and temperatures in the range 15–25 °C was used to estimate the lower temperature threshold for oviposition as 11.4 °C. With a nonlinear model fitted to the data across the whole range of experimental temperatures, the lower and upper limiting temperatures and optimum temperature were found to be 7.9, 33.7, and 28.9 °C, respectively. A model for daily oviposition rate was fitted, which describes the pattern of oviposition over the entire oviposition period. 4. The analysis of life tables, combining developmental rates, reproduction, mortality, and sex ratio, suggests maximum population growth (r_m) at near 30 °C. 5. After generating a first brood, spruce bark beetles often re-emerge from the tree and produce other sister broods. The effects of temperature and number of sister broods on demography were evaluated using age-specific life-table analyses. It is hypothesized that sister broods play an important role in regions where I. typographus is monovoltine, but have only moderate significance where this species has more than one generation per season.     

Comparative multilocus phylogeography of two Palaearctic spruce bark beetles: influence of contrasting ecological strategies on genetic variation

While phylogeographic patterns of organisms are often interpreted through past environmental disturbances, mediated by climate changes, and geographic barriers, they may also be strongly influenced by species‐specific traits. To investigate the impact of such traits, we focused on two Eurasian spruce bark beetles that share a similar geographic distribution, but differ in their ecology and reproduction. Ips typographus is an aggressive tree‐killing species characterized by strong dispersal, whereas Dendroctonus micans is a discrete inbreeding species (sib mating is the rule), parasite of living trees and a poor disperser. We compared genetic variation between the two species over both beetles’ entire range in Eurasia with five independent gene fragments, to evaluate whether their intrinsic differences could have an influence over their phylogeographic patterns. We highlighted widely divergent patterns of genetic variation for the two species and argue that the difference is indeed largely compatible with their contrasting dispersal strategies and modes of reproduction. In addition, genetic structure in I. typographus divides European populations in a northern and a southern group, as was previously observed for its host plant, and suggests past allopatric divergence. A long divergence time was estimated between East Asian and other populations of both species, indicating their long‐standing presence in Eurasia, prior to the last glacial maximum. Finally, the strong population structure observed in D. micans for the mitochondrial locus provides insights into the recent colonization history of this species, from its native European range to regions where it was recently introduced.     

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.     

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

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Recapture of Ips typographus L. (Col., Scolytidae) with attractants of low release rates: localized dispersion and environmental influences

1 The dispersal of Ips typographus L. (Col., Scolytidae) was studied using a mark–release–recapture approach in a grid of traps equipped with pheromone lures of release rates of about 8.4 mg/day of 2‐methyl‐3‐buten‐2‐ol (MB) and 0.29 mg/day of (S)‐cis‐verbenol (cV) in experiment 1, and 1.2 mg/day of MB and 0.04 mg/day of cV in experiment 2. 2 We investigated whether beetle dispersal reflected the simple diffusion pattern observed in previous I. typographus experiments, for which attractant release rates generally approached 50 mg/day of MB and 1 mg/day of cV. We also examined how environmental parameters (wind) and human activities (felling) could influence the beetles' flight. 3 The recapture percentage was higher in experiment 1 than in experiment 2: respectively, 7.0% (with 64 traps) and 2.3% (with 100 traps) of the beetles that took off were caught in the traps. 4 With the higher release rate (experiment 1), trap catches decreased with increased distance, whereas with the lower release rate (experiment 2), trap catches rose between 50 and 100 m then decreased with increasing distance. 5 Flight was little orientated by prevailing wind directions, a feature probably explained by the low wind speeds (0–1.2 m/s) observed throughout the study. 6 High trap catches of unmarked beetles close to areas undergoing thinning activities suggest that the presence of freshly cut spruce and larch material could have an influence on dispersal, attracting the beetles into the felling area. Spatial analyses show that capture patterns were autocorrelated up to distances of about 250 m.     

Monitoring of Ips typographus and Pityogenes chalcographus: influence of trapping site and surrounding landscape on catches

• 1 The influence of trapping site (i.e. fresh clear‐cuts) characteristics and habitat amount (i.e. area of stands that may hold breeding material) in the surrounding landscape on catches of the bark beetles Ips typographus (L.) and Pityogenes chalcographus (L.) in pheromone‐baited flight‐barrier monitoring traps was studied.
• 2 For the two species, the study investigated: (i) the extent of the variation in catches among trapping sites; (ii) the extent of this variation that can be explained by models including trapping site characteristics (clear‐cut size, percentage of spruce in the cut stand, altitude) and habitat amounts in the surrounding landscape; and (iii) the spatial scale at which beetles respond to the habitat amount in the landscape.
• 3 The variation in catches among trap sites was 15‐fold larger for I. typographus than for P. chalcographus. There was a positive relationship between the catches of I. typographus and (i) the percentage of the surrounding landscape covered by mature spruce forest at radii 500–4000 m and (ii) the percentage of spruce in stands cut when the clear‐cuts used as trapping sites were created. For P. chalcographus, only the second relationship could be demonstrated.
• 4 Thus, for monitoring of I. typographus: (i) several trapping sites per landscape are required; (ii) the amount of mature spruce forest around trapping sites needs to be considered when choosing trapping sites; and (iii) the trapping sites need to be standardized with respect to the percentage of spruce in the cut stand when fresh clear‐cuts are used. For P. chalcographus, fewer trapping sites per landscape are required and only the percentage of spruce in the cut stand needs to be considered.

     

Temperature‐dependent development of the double‐spined spruce bark beetle Ips duplicatus (Sahlberg, 1836) (Coleoptera; Curculionidae)

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Fungi Vectored by the Bark Beetle Ips typographus Following Hibernation Under the Bark of Standing Trees and in the Forest Litter

The bark beetle Ips typographus has different hibernation environments, under the bark of standing trees or in the forest litter, which is likely to affect the beetle-associated fungal flora. We isolated fungi from beetles, standing I. typographus-attacked trees, and forest litter below the attacked trees. Fungal identification was done using cultural and molecular methods. The results of the two methods in detecting fungal species were compared. Fungal communities associated with I. typographus differed considerably depending on the hibernation environment. In addition to seven taxa of known ophiostomoid I. typographus-associated fungi, we detected 18 ascomycetes and anamorphic fungi, five wood-decaying basidomycetes, 11 yeasts, and four zygomycetes. Of those, 14 fungal taxa were detected exclusively from beetles that hibernated under bark, and six taxa were detected exclusively from beetles hibernating in forest litter. The spruce pathogen, Ceratocystis polonica, was detected occasionally in bark, while another spruce pathogen, Grosmannia europhioides, was detected more often from beetles hibernating under the bark as compared to litter. The identification method had a significant impact on which taxa were detected. Rapidly growing fungal taxa, e.g. Penicillium, Trichoderma, and Ophiostoma, dominated pure culture isolations; while yeasts dominated the communities detected using molecular methods. The study also demonstrated low frequencies of tree pathogenic fungi carried by I. typographus during its outbreaks and that the beetle does not require them to successfully attack and kill trees.     

Can observation of climatic variables be used to predict the flight dispersal rates of Prostephanus truncatus?

Abstract 1 Attack by Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae) on the maize and cassava stored by small‐holder farmers in Africa is sporadic, varying considerably within and between years. The risk that food in store will become infested is related to the number of beetles dispersing by flight. A means of predicting years with high dispersal rates is needed to warn farmers when to be vigilant. 2 The relationship between climatic variables and pheromone trap catches was observed in a forest‐savannah transition zone in Ghana. These observations were used to devise a model using a mix of biological and empirical rules that operate on temperature and humidity data. The predicted and actual trap catch deviated by only +5% to ?1% in years when there were high dispersal rates. 3 The first part of the model estimates the numbers of beetles with potential for dispersal. The second part predicts the proportion likely to disperse. This is based on the apparent effect that those P. truncatus developing under low temperature conditions (about 24 °C) have a lowered propensity for flight, a response previously observed in a related species. 4 The model was validated using climate data and trap catches from a woodland–savannah zone and a short grass steppe zone. With minor adjustment, the model worked well for these two habitats.     

Exogenous application of methyl jasmonate elicits defenses in Norway spruce (Picea abies) and reduces host colonization by the bark beetle Ips typographus

The terpenoid and phenolic constituents of conifers have been implicated in protecting trees from infestation by bark beetles and phytopathogenic fungi, but it has been difficult to prove these defensive roles under natural conditions. We used methyl jasmonate, a well-known inducer of plant defense responses, to manipulate the biochemistry and anatomy of mature Picea abies (Norway spruce) trees and to test their resistance to attack by Ips typographus (the spruce bark beetle). Bark sections of P. abies treated with methyl jasmonate had significantly less I. typographus colonization than bark sections in the controls and exhibited shorter parental galleries and fewer eggs had been deposited. The numbers of beetles that emerged and mean dry weight per beetle were also significantly lower in methyl jasmonate-treated bark. In addition, fewer beetles were attracted to conspecifics tunneling in methyl jasmonate-treated bark. Stem sections of P. abies treated with methyl jasmonate had an increased number of traumatic resin ducts and a higher concentration of terpenes than untreated sections, whereas the concentration of soluble phenolics did not differ between treatments. The increased amount of terpenoid resin present in methyl jasmonate-treated bark could be directly responsible for the observed decrease in I. typographus colonization and reproduction.     

Temperature Determines Symbiont Abundance in a Multipartite Bark Beetle-fungus Ectosymbiosis

In this study, we report evidence that temperature plays a key role in determining the relative abundance of two mutualistic fungi associated with an economically and ecologically important bark beetle, Dendroctonus ponderosae. The symbiotic fungi possess different optimal temperature ranges. These differences determine which fungus is vectored by dispersing host beetles as temperatures fluctuate over a season. Grosmannia clavigera is the predominant fungus carried by dispersing beetles during cool periods but decreases in prevalence as daily maximum temperatures approach 25°C, and becomes extremely rare when temperatures reach or exceed 32°C. In contrast, Ophiostoma montium increases in prevalence as temperatures approach 25°C, and becomes the predominant symbiont dispersed when temperatures reach or exceed 32°C. The possession of different optimal growth temperatures may facilitate the stable coexistence of the two fungi by supporting growth of each fungus at different times, minimizing direct competition. Furthermore, the beetle may reduce its risk of being left aposymbiotic by exploiting not one, but two symbionts, whose combined growth optima span a wide range of environmental conditions. The possession of multiple symbionts with different temperature tolerances may allow the beetle to occupy highly variable habitats over a wide geographic range. Such temperature-driven symbiont shifts are likely to have major consequences for both the host and its symbionts under current temperature regimes and those predicted to occur because of climate change.     

Functional investigation of monoterpenes for improved understanding of the relationship between hosts and bark beetles

Spruce bark beetle (Ips typographus L.) is the most destructive insect pest of spruce forests in Eurasia. However, contact toxicity, in vivo metabolism, and ecological functions of host monoterpenes are poorly understood at the spruce tree–bark beetle–predator tritrophic level. Spruce monoterpenes including S-(–)-α-pinene, R-(+)-α-pinene, and myrcene showed contact toxicity to I. typographus, with LD₅₀ values ranging from 22–32 μg/mg. When topically treated with S-(–)-α-pinene or R-(+)-α-pinene, the amounts of volatile metabolites, including 4S-(–)-cis-verbenol, 4S-(+)-/4R-(–)-trans-verbenol, R-(+)-/S-(–)-verbenone and 1R-(–)-/1S-(+)-myrtenol, in the hindgut extracts of I. typographus varied significantly between sexes, and their quality (enantiomeric composition) varied significantly with the chirality of α-pinene. More importantly, S-(–)-α-pinene induced male adults to produce large amounts of 4S-(–)-cis-verbenol and S-(–)-verbenone. When topically treated with myrcene, the expected semiochemicals such as E-myrcenol, ipsenol and ipsdienol were not detected in the beetle hindguts, indicating that the pheromone biosynthetic system of I. typographus does not participate in the metabolism of host myrcene. In trap tests, S-(–)-α-pinene and R-(+)-α-pinene increased the catches of I. typographus and its predator Thanasimus substriatus in pheromone-baited traps, whereas myrcene exhibited a strong repellent (or inhibitory) effect on I. typographus but not on its predator. I. typographus seems to adopt different ecological strategies (e.g. avoidance to myrcene and preference for α-pinene) to adapt to and tolerate different host monoterpenes. Extensive investigation of these monoterpenes will help us understand their roles in manipulating the arms race between host trees and bark beetles, and potentially improve the efficacy of controlling I. typographus via the push-pull strategy using host kairomones.     

The neotropical shrub Lupinus elegans,from temperate forests,may not adapt to climate change

Considering that their distribution is limited to altitudinal gradients along mountains that are likely to become warmer and drier, climate change poses an increased threat to temperate forest species from tropical regions. We studied whether the understorey shrub Lupinus elegans, endemic to temperate forests of west‐central Mexico, will be able to withstand the projected temperature increase under seven climate change scenarios. Seeds were collected along an altitudinal gradient and grown in a shade‐house over 7 months before determining their temperature tolerance as electrolyte leakage. The plants from colder sites tolerated lower temperatures, i.e. the temperature at which half of the maximum electrolyte leakage occurred (LT₅₀), ranged from −6.4 ± 0.7 to −2.4 ± 0.3 °C. In contrast, no pattern was found for tolerance to high temperature (LT₅₀ average 42.8 ± 0.3 °C). The climate change scenarios considered here consistently estimated an increase in air temperature during the present century that was higher for the maximum air temperature than for the mean or minimum. In particular, the anomaly from the normal maximum air temperature at the study region ranged from 2.8 °C by 2030 to 5.8 °C by 2090. In this respect, the inability of L. elegans to adapt to increasingly higher temperatures found here, in addition to a possible inhibition of reproduction caused by warmer winters, may limit its future distribution.     

Pathogens of the spruce bark beetles Ips typographus and Ips duplicatus

Pathogens of two important bark beetles, Ips typographus and Ips duplicatus, both in outbreaks connected with infestation of spruces by the fungus Armillaria ostoyae, were compared at four localities in the eastern Czech Republic. Low infestations of Chytridiopsis typographi, Nosema typographi, Menzbieria chalcographi, and Gregarina typographi were detected in I. typographus. In I. duplicatus, only C. typographi and G. typographi were found and with low infection levels. The microsporidium, Larssoniella duplicati, was not detected in I. typographus, but was detected in I. duplicatus at all localities in almost 80% of the samples (a sample consisted of 40–50 beetles collected at one locality in one period) and often with a very high infection level (up to 57% of the beetles infected in a sample). The infection level of L. duplicati did not differ between generations of I. duplicatus. I. duplicatus overwinters mainly in the adult stage, and no decrease in the number of infected overwintering I. duplicatus was observed. The relatively constant infection level of L. duplicati suggests that transmission is unlikely to be horizontal via oral ingestion.     

A host monoterpene influences Ips typographus (Coleoptera: Curculionidae,Scolytinae) responses to its aggregation pheromone

1 Host tree terpenes can influence attraction of conifer‐infesting bark beetles to their aggregation pheromones, and both synergistic and inhibitory effects have been reported. 2 We tested a gradient of ratios of (–)‐α‐pinene, the predominant monoterpene in Norway spruce, to the pheromone of Ips typographus, a major pest of Norway spruce. 3 Attraction of I. typographus increased as the release rate of (–)‐α‐pinene increased. The two highest (–)‐α‐pinene : pheromone ratios (526 : 1 and 2595 : 1) attracted twice as many I. typographus as pheromone alone, whereas low to intermediate ratios (56 : 1, 274 : 1) did not differ from pheromone alone. 4 Our results are in agreement with a proposed model, which suggests that bark beetles display unique response profiles to host terpenes depending on the physiological condition of the host trees that they typically colonize. Ips typographus, which is an aggressive species capable of colonizing and killing healthy trees, showed an increased attraction to monoterpene : pheromone ratios, and this may be high enough to inhibit attraction of less aggressive beetle species typically colonizing dead, dying or stressed trees. 5 Attraction of associates of I. typographus was also modified by (–)‐α‐pinene. Ips duplicatus, a competitor of I. typographus, showed increased attraction to the pheromone of I. typographus across all concentrations of (–)‐α‐pinene.     

Pathogens of Ips amitinus: new species and comparison with Ips typographus

Ips amitinus and I. typographus are two serious pests of spruce in Europe, have similar bionomics and are likely to occur and meet on the same host trees. We therefore hypothesized that the two species support similar levels of similar pathogens. To test this hypothesis, we collected mature beetles from three trap trees at each of eight study sites and determined beetle numbers and pathogen infection levels. In total, 938 mature I. amitinus beetles and 3435 of I. typographus were dissected; five pathogens, as well as intestinal nematodes and endoparasitoids, were detected. The neogregarine Mattesia schwenkei is reported here for the first time as a new pathogen in 9.4% of I. amitinus individuals at one site. Average infection levels of most pathogens (Chytridiopsis typographi, Gregarina typographi, Mattesia schwenkei and parasitoids) were significantly higher in I. typographus than in I. amitinus. Metschnikowia typographi was confirmed only in Ips amitinus, while the microsporidium of Nosema typographi occurred only in I. typographus. Within‐season increases in G. typographi infection levels were documented in Ips amitinus.     

Divergent evolutionary histories of two sympatric spruce bark beetle species

Ips typographus and Pityogenes chalcographus are two sympatric Palearctic bark beetle species with wide distribution ranges. As both species are comparable in biology, life history, and habitat, including sharing the same host, Picea abies, they provide excellent models for applying a comparative approach in which to identify common historical patterns of population differentiation and the influence of species-specific ecological characteristics. We analysed patterns of genetic diversity, genetic structure and demographic history of ten I. typographus and P. chalcographus populations co-distributed across Europe using both COI and ITS2 markers. Rather than similarities, our results revealed striking differences. Ips typographus was characterised by low genetic diversity, shallow population structure and strong evidence that all extant haplogroups arose via a single Holocene population expansion event. In contrast, genetic variation and structuring were high in P. chalcographus indicating a longer and more complex evolutionary history. This was estimated to be five times older than I. typographus, beginning during the last Pleistocene glacial maximum over 100 000 years ago. Although the expansions of P. chalcographus haplogroups also date to the Holocene or just prior to its onset, we show that these occurred from at least three geographically separated glacial refugia. Overall, these results suggest that the much longer evolutionary history of P. chalcographus greatly influenced the levels of phylogeographic subdivision among lineages and may have led to the evolution of different life-history traits which in turn have affected genetic structure and resulted in an advantage over the more aggressive I. typographus.     

Water and energy fluxes during summer in an arid‐zone passerine bird

Endothermic animals resident in hot, arid terrestrial environments are likely to face a trade‐off between their ability to obtain water and elevated thermoregulatory water requirements. We assessed whether daily water flux (DWF) is higher on hot days, reflecting increases in evaporative cooling demands, in an arid‐zone bird that obtains its water through food intake. We obtained measurements of DWF (partitioned into water influx and efflux rates) in 71 White‐browed Sparrow‐Weavers Plocepasser mahali at a desert site and a semi‐desert site, during summer in the Kalahari Desert of southern Africa. We found no evidence that DWF varied with maximum daily air temperature (T_air, range = 27.6–39.2 °C). Instead, DWF was lower during dry periods than in the wet season at the semi‐desert site. Furthermore, birds showed deficits in water balance (water influx/water efflux) during the dry periods at both sites. Our data show that DWF is low in a non‐drinking bird that obtains its water through food, and that demands for evaporative water loss on very hot days (maximum T_air of 40–44 °C) may exceed water intake rates during hot and dry periods. Species that do not have opportunities to drink will experience strong trade‐offs between thermoregulation, hydration state and activity levels as temperatures increase.     

Biology and associated fungi of an emerging bark beetle pest,the sweetgum inscriber Acanthotomicus suncei (Coleoptera: Curculionidae)

The sweetgum inscriber (SI) Acanthotomicus suncei Cognato is an emerging bark beetle pest that seriously damages American sweetgum trees (Liquidambar styraciflua) and Chinese sweetgum trees (L. formosana) in China. Since 2013, SI has killed more than 30,000 sweetgum trees in Shanghai and adjacent areas. In Shanghai, SI was observed to emerge from infested wood between April–September and exhibited two generations per year. Both the flying populations and those in colonized logs were female-biased. After the parental adult initiates a gallery, the emergence of new adults required 73.4 ± 0.7 days at 30°C and 163.5 ± 0.9 days at 20°C. SI could not sustain population development at constant temperatures ≤15°C. The fungi Geosmithia spp. and Phaeoacremonium tuscanum were common associates of SI. The results suggest that SI has a long flight period, and its development is sensitive to low temperatures. Fungi associated with SI were not pathogenic to American sweetgum trees, but their presence may indirectly increase the probability of tree mortality. Our findings contribute to the prevention and control of this important emerging bark beetle pest.