Evidence that the availability of suitable pine limits non‐native Sirex noctilio in Ontario

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Emergence of adult female Sirex nigricornis F. and Sirex noctilio F. (Hymenoptera: Siricidae) coincides with a decrease in daily minimum and maximum temperature

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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.

     

A comparison of control results for the alien invasive woodwasp,Sirex noctilio,in the southern hemisphere

1 Sirex noctilio has resulted in one of the most damaging biological invasions of pine forestry in the southern hemisphere. 2 An intensive, integrated control programme has been developed for this pest and is generally considered very successful. However, a critical comparison of S. noctilio infestations and control efforts throughout the southern hemisphere reveals that control has not been uniformly effective. Of particular concern is the apparently unexplained variability in establishment and success of biological control agents, including various parasitic wasps and the parasitic nematode Deladenus siricidicola. 3 An overlooked aspect in the establishment of the biological control interventions for S. noctilio is the potential influence of genetic bottlenecks on the ability of the agents to adapt to different environments and different populations of S. noctilio. 4 Despite established biological control agents, stress in forests caused by silvicultural practices or the environment will predispose trees to heavy infestation. Unfortunately, improved silvicultural practices are not always economically feasible and environmental changes are often unavoidable. 5 Sirex noctilio continues to pose a serious threat to pine forestry in many areas. Despite extensive earlier research into a single integrated control for S. noctilio, it is important to recognize that such a strategy will probably require significant local adaptation in new areas of infestation and over time.     

Symbionts mediate oviposition behaviour in invasive and native woodwasps

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Local antagonism and resource partitioning between two invasive pine plantation pests

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Sirex noctilio in North America: the effect of stem‐injection timing on the attractiveness and suitability of trap trees

• 1 Sirex noctilio Fabricius, an invasive woodwasp responsible for severe economic damage to pine industries in the southern hemisphere, is now established in the northeastern U.S.A. and portions of eastern Canada.
• 2 Parts of North America are considered to be high risk for S. noctilio invasion. Effective detection tools, including trap trees, are needed to monitor and survey S. noctilio populations.
• 3 The present study was conducted to determine the optimal time to chemically stress a tree when aiming to attract the most S. noctilio to the host substrate, as well as to determine which timing produced the most adult progeny. Both of these measures (host attraction and host suitability for development) support the main objectives of the study by offering improved methods for monitoring and management of S. noctilio.
• 4 Red pine (Pinus resinosa) and Scots pine (Pinus sylvestris) were treated with Dicamba at three time intervals. Multiple funnel lindgren traps were placed on these trees and, at the end of the flight season, the treatment trees were felled and brought into the laboratory. The number of S. noctilio caught in the traps (host attraction) and the number of S. noctilio emerged from the treated trees (host suitability) were determined.
• 5 Optimal timing of the chemical girdle was dependent on host species. Significantly more female S. noctilio were captured on trap trees prepared 1 month before flight (red pine and Scots pine) or prepared at flight (Scots pine) compared with other treatments. There were also significantly more females reared from Scots pine trap trees prepared at flight and red pine trap trees prepared 1 month before and/or at flight.
• 6 By the beginning of August, most (79%) of the S. noctilio for the flight season were caught in the traps at the trap trees. The sex ratio (males : females) was closer to 1 : 1 than previously reported in studies from other countries.
• 7 The results obtained in the present study demonstrate that timing is important when creating a trap tree with herbicide in North America, whether for the purpose of detection or as part of a biological control effort.

     

Assessing trap and lure effectiveness for the monitoring of Sirex noctilio

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Phenology and flight periodicity of Sirex noctilio (Hymenoptera: Siricidae) in central New York,U.S.A.

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The importance of olfactory and visual cues in developing better monitoring tools for Sirex noctilio (Hymenoptera: Siricidae)

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Mechanisms that influence sex ratio variation in the invasive hymenopteran Sirex noctilio in South Africa

Sirex noctilio is an economically important invasive pest of commercial pine forestry in the Southern Hemisphere. Newly established invasive populations of this woodwasp are characterized by highly male‐biased sex ratios that subsequently revert to those seen in the native range. This trend was not observed in the population of S. noctilio from the summer rainfall regions in South Africa, which remained highly male‐biased for almost a decade. The aim of this study was to determine the cause of this persistent male bias. As an explanation for this pattern, we test hypotheses related to mating success, female investment in male versus female offspring, and genetic diversity affecting diploid male production due to complementary sex determination. We found that 61% of females in a newly established S. noctilio population were mated. Microsatellite data analysis showed that populations of S. noctilio from the summer rainfall regions in South Africa are far less genetically diverse than those from the winter rainfall region, with mean Nei's unbiased gene diversity indexes of 0.056 and 0.273, respectively. These data also identified diploid males at low frequencies in both the winter (5%) and summer (2%) rainfall regions. The results suggest the presence of a complementary sex determination mechanism in S. noctilio, but imply that reduced genetic diversity is not the main driver of the male bias observed in the summer rainfall region. Among all the factors considered, selective investment in sons appears to have the most significant influence on male bias in S. noctilio populations. Why this investment remains different in frontier or early invasive populations is not clear but could be influenced by females laying unfertilized eggs to avoid diploid male production in populations with a high genetic relatedness.     

Host tissue identification for cryptic hymenopteran parasitoids associated with Sirex noctilio

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Effect of temperature on woodwasp (Sirex noctilio F.) development and parasitism by the entomopathogenic nematode,Deladenus siricidicola

The woodwasp, Sirex noctilio, is a significant global pest of exotic pine plantations in the Southern Hemisphere and now threatens native pine forests in North America. Management in Australia relies on biocontrol using the nematode, Deladenus (= Beddingia) siricidicola (Bedding), which infects and sterilises females who then further disperse the nematode. This pest is spreading into warmer regions in Australia and South America and coupled with the threat of global climate change, there is uncertainty as to how increasing temperatures will affect the biocontrol program. S. noctilio within nematode-inoculated wood were reared at four temperatures (24, 25.3, 26.6 and 28 °C) to investigate the effects of elevated temperatures on wasp development (emergence time, sex ratio and size), development of eggs (number, size, and maturation) and infection by the nematode. At 24 °C, which reflects current field temperature, S. noctilio were bigger in size and all the eggs were normal and all were infected with nematodes. Modest rises in temperature reflecting climate change scenarios resulted in smaller sized S. noctilio, disrupted egg development and maturation, and lowered the nematode sterilisation rate in females. Reduced S. noctilio female body size and egg infection will likely compromise biocontrol by D. siricidicola in its current distribution, but disrupted egg development may act directly on the pest, limiting dispersal of S. noctilio into subtropical pine plantations and adaptation to climate change.     

Spatial Pattern of Attacks of the Invasive Woodwasp Sirex noctilio,at Landscape and Stand Scales

Invasive insect pests are responsible for important damage to native and plantation forests, when population outbreaks occur. Understanding the spatial pattern of attacks by forest pest populations is essential to improve our understanding of insect population dynamics and for predicting attack risk by invasives or planning pest management strategies. The woodwasp Sirex noctilio is an invasive woodwasp that has become probably the most important pest of pine plantations in the Southern Hemisphere. Our aim was to study the spatial dynamics of S. noctilio populations in Southern Argentina. Specifically we describe: (1) the spatial patterns of S. noctilio outbreaks and their relation with environmental factors at a landscape scale; and (2) characterize the spatial pattern of attacked trees at the stand scale. We surveyed the spatial distribution of S. noctilio outbreaks in three pine plantation landscapes, and we assessed potential associations with topographic variables, habitat characteristics, and distance to other outbreaks. We also looked at the spatial distribution of attacked trees in 20 stands with different levels of infestation, and assessed the relationship of attacks with stand composition and management. We found that the spatial pattern of pine stands with S. noctilio outbreaks at the landscape scale is influenced mainly by the host species present, slope aspect, and distance to other outbreaks. At a stand scale, there is strong aggregation of attacked trees in stands with intermediate infestation levels, and the degree of attacks is influenced by host species and plantation management. We conclude that the pattern of S. noctilio damage at different spatial scales is influenced by a combination of both inherent population dynamics and the underlying patterns of environmental factors. Our results have important implications for the understanding and management of invasive insect outbreaks in forest systems.     

A double-edged sword: Amylostereum areolatum odors attract both Sirex noctilio (Hymenoptera: Siricidae) and its parasitoid,Ibalia leucospoides

The Eurasian woodwasp, Sirex noctilio, is an invasive pest of pines in temperate regions of the Southern Hemisphere, and has been introduced to North America. A parasitoid, Ibalia leucospoides, has been broadly employed for biological control of this pest. Volatiles emitted from the fungal symbiont of S. noctilio, Amylostereum areolatum, are reliable cues for S. noctilio and I. leucospoides females to optimize their foraging behavior (host location and host habitat finding) in a chemically complex environment. The headspace volatiles of A. areolatum, were analyzed using coupled gas chromatography-mass spectrometry (GC-MS) and GC-electroantennographic detection (GC-EAD) for both S. noctilio and I. leucospoides females. Analyses revealed that both species could detect several fungal volatiles. In olfactometer bioassays, S. noctilio females were attracted to a 4-component blend of 6-methyl-5-hepten-2-one, trans-3-hexenyl acetate, linalool, and geraniol, while the addition of ρ-anisaldehyde to the blend was necessary for attraction of I. leucospoides females. The results of trap catches in field experiments confirmed that these fungal volatiles in combination with host tree volatiles are attractive to both species, although the release rate of the fungal volatiles is important. These volatiles can serve as a basis for the development of improved lures for both species.     

Delayed emergence and the success of parasitoids in biological control

Delayed emergence, a life history feature of many insects living in unpredictable environments, can have major consequences for the dynamics of host–parasitoid interactions, which vary according to their physiological interactions. We studied, through simple modeling, the significance of prolonged diapause on the suppression levels achieved by parasitoids and illustrate our case with a system involving a major forest pest, the woodwasp Sirex noctilio and two of its parasitoid species that have been introduced into different geographical regions through classical biological control programmes. Our findings suggest that the physiological relationship between parasitoid and host delayed emergence patterns may help understand observed variable success in several bio-control programs. We conclude that for given environments, host delayed emergence and the way in which parasitoids deal with it, should be included in the list of selection criteria of natural enemies of many pests, especially those affecting forests.     

Trapping for Sirex Woodwasp in Brazilian Pine Plantations: Lure,Trap Type and Height of Deployment

The Sirex woodwasp, Sirex noctilio (Hymenoptera: Siricidae) is considered a secondary pest of pine in its native range but has caused considerable economic losses in pine plantation forests in the southern hemisphere. In Brazil, trap trees are the primary tool used for early detection purposes but these are costly, labor-intensive to install and require stressing trees by herbicide application. Flight intercept traps baited with synthetic blends of host volatiles are an attractive alternative but have performed poorly in some settings. This study was carried out to look for alternatives to trap trees for use in Brazilian pine plantations for early detection of S. noctilio. Four field experiments were conducted in two consecutive flight seasons (2015–16 and 2016–17), in planted loblolly pine (Pinus taeda) stands, to compare captures among flight intercept traps baited with different lures, deployed at different heights and among different intercept trap designs. Two experiments compared different host volatile lures and a significant treatment effect was observed in one. No effect of trap design or height was observed.     

Fidelity Among Sirex Woodwasps and Their Fungal Symbionts

We report that associations between mutualistic fungi and their economically and ecologically important woodwasp hosts are not always specific as was previously assumed. Woodwasps in the genus Sirex engage in obligate nutritional ectosymbioses with two species of Amylostereum, a homobasid\iomycete genus of white rot fungi. In the present study, the Amylostereum species and genotypes associated with three species of Sirex native to eastern North America and one relatively recent invasive Sirex from Europe were investigated by comparing intergenic spacer regions (IGS). Sirex spp. were sampled over 6 years from 23 sites in six US states, ranging from Maine in the northeast to Louisiana in the southeast, to obtain samples of Amylostereum from mycangia of adult females. Two of the native Sirex species (Sirex nigricornis and Sirex nitidus) were associated with either Amylostereum chailletii or Amylostereum areolatum, refuting the hypothesis of strict species-specific relationships. However, the invasive Sirex noctilio and the native Sirex cyaneus were each collected with only A. areolatum or A. chailletii, respectively, although S. noctilio was associated with two different IGS genotypes of A. areolatum and S. cyaneus occurs sympatrically with the other native Sirex. In Pinus, the preferred host tree of S. nigricornis and S. noctilio, these species co-occurred in 25.9 % of trees sampled, and horizontal transmission of fungal strains from S. noctilio to S. nigricornis was documented, although only in one tree. The extent that further spread and establishment of S. noctilio will alter the composition of symbionts carried by native Sirex is unknown but will depend in part on the degree of flexibility in these host–symbiont associations.     

Influence of the community of associates on Sirex noctilio brood production is contextual

1. Competition and predation are important components of biotic resistance, which helps define the invasibility of an ecosystem. 2. To search for evidence of biotic resistance to the European woodwasp, Sirex noctilio Fabricius, in North America, cages were used to experimentally exclude the community of associates (natural enemies and competitors) from infested logs. Specifically, the study assessed S. noctilio brood production in pine forests in Ontario and New York, where there was a rich existing community of associates (other wood borers, bark beetles and associated fungi, and parasitoids), and in South Africa, where siricid wasps and pines are not native and a similar associate community is not present. In addition, in Ontario, associates were excluded by size, and for different periods of time to identify important associates and their temporal dynamics. 3. Evidence was found that biotic factors limit S. noctilio in North America, whereby exclusion of natural enemies and competitors had a positive influence on the abundance or presence of S. noctilio brood in Ontario and New York. This influence was absent in South Africa. 4. It is unclear which member(s) of the associated insect community in North America were most important in limiting S. noctilio brood production, although they probably acted quickly (< 2 weeks) following S. noctilio oviposition. 5. Further study is needed to determine whether associates have limited S. noctilio populations in pine forests throughout northeastern North America, and which specific natural enemies and/or competitors are important.     

Evaluation of the insect parasitoids of Sirex noctilio (Hymenoptera: Siricidae) in Tasmania

Summary Sampling of Sirex noctilio F. and the insect parasitoids released in a forest of Pinus radiata D. Don near Hobart, Tasmania, has shown that the rhyssines, Megarhyssa nortoni (Cresson) and Rhyssa persuasoria (L.) were mainly responsible for reducing the level of the population of S. noctilio between 1965 and 1974. Parasitic nematodes were not present during this period. Key factor analysis indicates that in this forest the two species act as one delayed density-dependent factor, whereas Ibalia leucospoides Hochenw. is apparently density-independent, its ability to parasitize the host being regulated by conditions within the tree.