Auswahl der wissenschaftlichen Literatur zum Thema „Bark beetles Control“

Bark beetles (Coleoptera: Curculionidae) are among the important pests of forest in the world and in Turkey. Pine-feeding bark beetles interact chemically with their host tree via the behavioral, physiological, and biochemical effects of one class of monoterpenes. The half of oleoresin producing by trees such as pines, spruces, firs in the forests is the monoterpene. The monoterpenes occur in the oleoresin and function as behaviorally active kairomones for pine bark beetles and their predators, presenting a classic example of tritrophic chemical communication. The monoterpenes are aggregation pheromones for pine bark beetle. In the control of bark beetles, monoterpenes are used in traps. For example, the monoterpenes are an attractant for bark beetles. Also, Myrcene is used for a synergistic effect in the trap. Monoterpenes can toxic to insects. Limonene is among the most toxic monoterpenes to bark beetles. Also, the interaction is found between monoterpenes and prey of bark beetle. Monoterpenes are enhanced to respond of predatory to the pheromone of bark beetle.

AbstractThree field tests were conducted in which fresh lodgepole pine (Pinus contorta Douglas var. latifolia Engl.) material, namely bolts with and without bark, bark only, and freshly tapped resin, were placed in beetle-excluding “greenhouse” cages; empty cages served as controls. Two “window” flight traps per cage, at right angles to each other, caught mountain pine beetles (Dendroctonus ponderosae Hopkins) arriving at the cages. Significantly more mountain pine beetles were trapped at cages baited with bolts and wood only than at empty control cages. Primary attraction in the mountain pine beetle is thus established, in the absence of pheromones and normal visual cues (tree stem silhouette). More beetles were trapped at cages baited with bark only and with resin than at empty control cages, but differences were not significant at p = 0.05. The sex ratio of trapped beetles (4.83 females: 1 male) was more than twice as high as the reported sex ratios of free-flying and emerging beetles.

AbstractAlmost 100 years have passed since J.M. Swaine, the assistant entomologist in charge of Forest Insect Investigations, wrote, “Canadian bark-beetles: a preliminary classification, with an account of the habits and means of control”. The goal was to “put into the hands of practical foresters information of inestimable practical value… to prevent the continued loss of timber now being destroyed” by “the most insidious enemies of the forest”. In this paper, we celebrate Swaine’s pioneering work by summarising the foundational aspects of his early treatise of 1918: the “general habits” of bark beetles, classifications of their behaviour, causes of population increase, and mitigation tactics. In the founding text, Swaine identified all majorDendroctonusErichson (Coleoptera: Curculionidae) bark beetles found in Canada, although details on life histories were scarce. We summarise current knowledge of the life histories and population dynamics of the spruce beetle,D.rufipennisKirby; the Douglas-fir beetle,D. pseudotsugaeHopkins; the eastern larch beetle,D. simplex; and address the current range expansion of mountain pine beetle,D. ponderosaeHopkins. We review how aspects of global change, such as invasive species, have altered the population dynamics of certain bark beetles. Finally, we conclude with lessons from two of the many past contributors to bark beetle ecology in Canada, J.M. Swaine and H.A. Richmond.

Two experiments were conducted in mixed hardwood-conifer forests in the northeastern United States to test the effects of cleaning surfactant and non-surfactant treated multiple-funnel traps used to catch bark and woodboring beetles. Large amounts of pollen and other debris often form a crust on the interior of traps (personal observations). Such surface deposits may provide footholds for beetles to escape capture in traps. In one experiment, we tested cleaned surfactant and non-surfactant traps against non-cleaned surfactant and non-surfactant traps. In a second experiment, we tested field cleaning of modified multiple-funnel traps as an alternative to substituting clean traps on each collection visit. There was no effect of surfactant treated traps, cleaned or not, on total beetles or individual bark beetle species captured. However, in situ cleaned traps were statistically better at capturing total beetles, total bark beetles, and several bark beetle species than non-cleaned control traps. Surfactant-treated non-modified traps and cleaned modified traps had higher species richness and abundance than other treatments at the site level. Our results suggest that cleaning traps to remove accumulated pollen and debris may be helpful for some species but would have limited benefit for broad-scale trapping of bark and woodboring beetles in northeastern forests.

The success of tree colonization by bark beetles depends on their ability to overcome host tree defenses, including resin exudation and toxic chemicals, which deter bark beetle colonization. Resin defenses during insect outbreaks are challenging to study in situ, as outbreaks are stochastic events that progress quickly and thus preclude the establishment of baseline observations of non-infested controls. We use synthetic aggregation pheromones to demonstrate that confined Ips bark beetle herbivory can be successfully initiated to provide opportunities for studying interactions between bark beetles and their hosts, including the dynamics of constitutive and induced resin exudation. In Pinus taeda L. plantations between 12 and 19 years old in North and South Carolina, U.S., trees were affixed with pheromone lures, monitored for evidence of bark beetle attacks, and resin samples were collected throughout the growing season. Baiting increased beetle herbivory to an extent sufficient to produce an induced resin response. Attacked trees exuded about three times more resin at some time than control trees. This supports previous work that demonstrated that information on constitutive resin dynamics alone provides an incomplete view of a host tree’s resistance to bark beetle attack.

Abstract The study estimated the efficacy of a net coated with alpha-cypermethrin used to protect Norway spruce [Picea abies (L.) H. Karst] wood against bark beetle infestations. The inside of the net was coated with 100 mg/m2 of alpha-cypermethrin. The studies carried out in 2010 and 2011 took place in southern Poland in the Beskid Żywiecki and Beskid Sądecki mountains in P. abies stands threatened by Ips typographus (Linnaeus, 1758). The research material consisted of wood logs taken off of 70–80-year-old P. abies trees. The logs were wrapped in the net in the early spring before spring swarming of bark beetles. The unwrapped logs were considered as the study controls. An evaluation of the treatments was performed after 2 months. The evaluation was based on the counting of bark beetles galleries found after removing the bark from the wrapped and unwrapped logs. There were no insect galleries on the wrapped logs. The net was a barrier, on which bark beetles died. About 10 dead I. typographus beetles were found on 0.01 m2 of the net surface. On the bark from the unwrapped-control logs there were 3,156 galleries/entrance holes of bark beetles, in total, of which 73% belonged to I. typographus, 13% to Pityogenes chalcographus (Linnaeus, 1761), and almost 10% to Xyloterus lineatus (Oliv.). These results indicated the high efficacy of the net coated with alpha-cypermethrin used for the protection of Norway spruce wood against the bark beetles. At the same time, the net was found to have a negative effect on non-target entomofauna, mainly Hylobius spp., Thanasimus formicarius (L.), and Tetropium castaneum (F.). However, the use of a net did not affect other predatory (Carabidae) and parasitic (Ichneumonidae and Tachinidae) entomofauna.

Bark beetles are among the most influential biotic agents in conifer forests, and forest management often focuses on bark beetle chemical communication for tree protection. Although acoustic communication occurs in many bark beetle species, we have yet to utilize acoustic communication for bark beetle control. Here, we describe the stridulatory organs and ‘stress’ chirps of the pinyon engraver, Ips confusus, a significant pest and mortality agent of pinyon pine in western North America. Only females possessed stridulatory organs and their stress chirps varied significantly in duration, pulses per chirp, and dominant frequency. We tested an array of acoustic-vibrational treatments into logs but were unable to disrupt male entry into logs or alter female–male interactions, female tunneling, and female oviposition. We found acoustic–vibrational treatments had little effect on I. confusus behavior and suggest further studies if acoustic methods are to be utilized for bark beetle control.

Research Highlights: Bark beetles are important agents of disturbance regimes in temperate forests, and specifically in a connected wind-bark beetle disturbance system. Large-scale windthrows trigger population growth of the European spruce bark beetle (Ips typographus L.) from endemic to epidemic levels, thereby allowing the killing of Norway spruce trees over several consecutive years. Background and Objectives: There is a lack of evidence to differentiate how outbreaks are promoted by the effects of environmental variables versus beetle preferences of trees from endemic to outbreak. However, little is known about how individual downed-tree characteristics and local conditions such as tree orientation and solar radiation affect beetle colonization of downed trees. Materials and Methods: To answer this question, we investigated the infestation rates and determined tree death categories (uprooted, broken, and stump) in wind-damaged areas in Western Tatra Mts. in Carpathians (Slovakia) from 2014–2016, following a windthrow in May 2014. In total, we investigated 225 trees over eight transects. For every tree, we measured its morphological (tree height, crown characteristics), environmental (solar radiation, terrain conditions, trunk zenith), temporal (time since wind damage), and beetle infestation (presence, location of attack, bark desiccation) parameters. We applied Generalized Additive Mixed Models (GAMM) to unravel the main drivers of I. typographus infestations. Results: Over the first year, beetles preferred to attack broken trees and sun-exposed trunk sides over uprooted trees; the infestation on shaded sides started in the second year along with the infestation of uprooted trees with lower desiccation rates. We found that time since wind damage, stem length, and incident solar radiation increased the probability of beetle infestation, although both solar radiation and trunk zenith exhibited nonlinear variability. Our novel variable trunk zenith appeared to be an important predictor of bark beetle infestation probability. We conclude that trunk zenith as a simple measure defining the position of downed trees over the terrain can anticipate beetle infestation. Conclusions: Our findings contribute to understanding of the bark beetle’s preferences to colonize windthrown trees in the initial years after the primary wind damage. Further, our findings can help to identify trees that are most susceptible to beetle infestation and to prioritize management actions to control beetle population while maintaining biodiversity.

Golovyanko Zinoviy Stepanovich is an outstanding entomologist well-known in Ukraine and abroad, specialist in insects, which are pests of forest. He is a Doctor of agricultural sciences, Professor. His scientific activity was generally devoted to working out of the measures on creating of pine plantations in the conditions of high number of the roots’ pests, which are largely grubs of May beetle (Melolontha melolontha L., M. hippocastani F.) and July beetle (Polyphillo fullo L.). He revealed that the density of May beetle’s grubs in the soil depends on temperature of this media. Не revealed the capacity of a pine to resist themselves against pests of their roots; this capacity was provided by oleoresin exudation from wounds, which were inflicted by the grubs on roots. He proposed recommendations on pine and deciduous tree species protection from May beetle’s grubs. He developed control measures against scarab beetles grubs for the first time. Z.S. Golovyanko brought the fundamental contribution into solution of the problem on revealing of the factors of population dynamics of bark beetles (Coleopera, Curculionidae: Scolytinae). He had worked out chemical method of forest pests’ control by use of aviation too. The scientific and practical heritage of Z.S. Golovyanko on ecological approach to forest protection from pest organisms is very actual in the present time.

Nematode parasites of bark beetles are taxonomically diverse, and may, in some cases, cause serious pathological and behavioural changes in their hosts (Kaya 1984). Their potential as biological control agents has long been recognized, but to date there is a shortage of elementary information on host-parasite interactions. Though the effects of superparasitism and subsequent crowding on the parasite have been studied for some mermithids (Petersen 1972) and steinernematids (Sandner and Stanuszek 1971), no such observations have been reported for bark beetle parasites. During a survey of nematode parasites of bark beetles in Manitoba, we encountered two examples where nematodes were detrimentally affected at the higher infection rates observed. It was our objective, therefore, to examine the relationship between intensity of infection, and impact on growth and reproduction of the parasites.

Nos recherches abordent, par une approche multidisciplinaire, l’écologie spatiale et la dynamique d’invasion du ravageur de l’épicéa Ips typographus (Coleoptera :Scolytinae) en Belgique. Bien que cette espèce ne puisse pas être considérée au sens strict comme invasive en Belgique où elle est implantée depuis le milieu du 19ème siècle, de très vraisemblables introductions de souches exotiques en provenance de Russie et des Pays baltes, en font un modèle exceptionnel pour l’étude de ces processus.

Une analyse en milieu urbain, suivant des transects centre-périphérie, a d’une part suggéré l’introduction probable de ces souches exotiques et d’autre part permis d’étudier ce ravageur forestier dans un milieu qui lui est a priori défavorable. Il en résulte que, malgré un effet de la structure urbaine sur l’abondance du scolyte, sa présence a été observée partout, y compris dans les zones densément bâties du centre de Bruxelles. Cela suggère, en terme de dispersion, que cette espèce est capable d’atteindre des arbres isolés, et en terme d’infestation, que tout épicéa constitue un hôte potentiel tant en milieu forestier qu’en milieu ouvert ou urbain.

Une étape préliminaire dans l’estimation des risques liés aux invasions biologiques est d’effectuer une analyse de filière, afin d’analyser les différentes voies d’introduction et les risques liés aux pays d’origine des marchandises importées. Sur base de l’analyse de l’ensemble des sources de données statistiques disponibles, nous avons déterminé l’ampleur du commerce de grumes d’épicéa entre l’est de l’Europe et la Belgique, les modes de transport utilisés, les origines et destinations des grumes, ainsi que la variabilité de la localisation et du nombre des différents points d’entrée de ces marchandises sur le territoire belge d’une année à l’autre, depuis le début des importations en 1996. Notre analyse illustre les difficultés inhérentes à ce genre d’étude, particulièrement au niveau de la disponibilité des données à une échelle appropriée.

Par une approche de modélisation, nous avons envisagé différents scénarios de dispersion de souches introduites d’Ips typographus afin d’estimer l’expansion de celles-ci en Belgique.

Notre modèle a permis de mettre en évidence l’importance de la bonne connaissance de certains paramètres biologiques caractétistiques de l’espèce étudiée (phénologie, capacité de dispersion, taux de croissance). Par ailleurs, les volumes d’importation et la localisation des points d’entrée des marchandises semblent jouer un rôle secondaire.

Enfin, nous avons utilisé des marqueurs génétiques afin d’essayer de prouver l’introduction de souches russes et baltes du scolyte de l’épicéa en Belgique. Des échantillons ont été collectés dans neuf pays européens ainsi que sur l’ensemble du territoire wallon. La structure génétique observée n’a pas permis de mettre en évidence une différenciation entre les populations de l’est et de l’ouest de l’Europe. La présence d’haplotypes communs à l’est et à l’ouest rejoint l’hypothèse d’une homogénéisation des populations en relation avec le large pouvoir de dispersion du typographe. Cependant, un grand nombre de mutations a été observé entre 2 haplotypes. Des introductions d’origines lointaines telles que la Sibérie ne sont donc pas improbables.

Les résultats détaillés de chacune de ces études sont développés dans les différents articles constituant cette thèse. La discussion générale met ceux-ci en relation et aborde les perspectives découlant de nos recherches.
Doctorat en sciences, Spécialisation géographie
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Bark beetles are major pests of pine forests in North America that can inflict considerable damage and cause severe economic loss. The checkered beetle Thanasimus dubius Fabricius (Coleoptera: Cleridae), an abundant predator in the Eastern USA and Canada, has been suggested to influence the dynamics of several bark beetles, including the economically important southern pine beetle (SPB), Dendroctonus frontalis Zimmermann (Coleoptera: Curculionidae). Because of the importance of T. dubius in pine forests, I studied several aspects of T. dubius biology including both its chemical and dispersal ecology. I first present an experiment that was conducted to improve the rearing system of T. dubius so that sufficient numbers of predators could be produced more efficiently for both behavioral and ecological studies. Improvements to the rearing method potentially could be achieved by adding a preservative to increase the shelf-life of the diet used to feed larval T. dubius thereby allowing feeding intervals to be extended. To evaluate this, I added sorbic acid, a preservative, to the larval diet at three different concentrations (0, 0.1 and 0.2%) and for two different intervals between feedings (2-3 vs. 5 days). Additionally, I measured potential deleterious effects of this preservative on predator performance (i.e., female fecundity). I then assessed the effect of this predator on SPB survival by releasing newly hatched T. dubius larvae at several densities (0, 50 100, and 200) on pine logs infested by SPB. I also studied the chemotactic response of this predator toward various volatiles such as prey pheromones or tree volatiles in a wind tunnel. I developed a method that associated upwind flight behavior and antennal behavior to quantify the relative attractiveness of each tested source of chemical. I then conducted three experiments to investigate the ability of individual T. dubius to respond to different prey olfactory signals (bark beetle pheromones) and assess their potential for learning. First, I examined individual predator preferences toward three major prey pheromones (ipsenol, ipsdienol, and frontalin) in a study that combined a mark-release-recapture experiment with a field choice test. I also measured the responses of individual wild T. dubius in a wind tunnel, which allowed the testing of a wider range of semiochemicals (frontalin, ipsenol, ipsdienol, sulcatol, and α-pinene). A final wind tunnel experiment tested the ability of naive predators to learn two prey pheromones, frontalin and ipsenol, that were presented alone or associated with a reward. In the last chapter, I quantified the dispersal of this predator and its behavior along an edge separating a pine forest from a non-suitable habitat. I analyzed T. dubius distribution and movement in the field relative to its bark beetle prey Ips grandicollis Eichhoff (Coleoptera: Curculionidae) and to the root feeder Dendroctonus valens LeConte (Coleoptera: Curculionidae). These three insects are associated with the phenomenon of red pine decline in the Great Lakes area. Thus, understanding the relative pattern of movements of these three beetles could provide considerable knowledge on the spatial and temporal progression of red pine decline by, for instance, assessing the connection between existing pockets. I estimated the dispersal quantiles for this predator relative to its bark beetle prey and determined whether dispersal behavior was relatively homogeneous (one kind of disperser) or heterogeneous (two kinds). In a second part, I sampled the abundance of T. dubius, I. grandicollis, and D. valens along transects set between a matrix area constituted by a clearing and a habitat zone composed of red pine forest. I further developed a diffusion model, including a constant k (corresponds to the ratio of densities along the edge), that permits characterization of beetle behavior around the boundary. Studies developed in the present dissertation, therefore, investigated several aspects of T. dubius ecology: predation on SPB, ability to respond to an olfactory signal, and dispersal traits. I first showed that augmentation with larval T. dubius can successfully reduce the SPB ratio of increase in infested pine logs. I also developed a system of rearing T. dubius that was more efficient and could allow larger numbers of predators to be produced. Sorbic acid did not reduce adult lifetime or size but did affect female fecundity (20 - 40% reduction). Increasing feeding time interval to 5 days (instead of 2-3 days) improved the efficiency of the system, even when sorbic acid was not added in the diet. Wind tunnel experiments present a methodology that could enable future quantification of the relative attractiveness of various semiochemical treatments. I identified a particular antennal behavior, which I called "stand up antennal" that was typically produced when the insect was attracted to a semiochemical. The choice tests in the field and the wind tunnel experiment using wild adults both showed that T. dubius individuals are generalists that are flexible in their response to a broad range of semiochemical signals (frontalin, ipsenol, ipsdienol, sulcatol, and α-pinene). I demonstrate that naive predators are more attracted toward ipsenol after having experienced conditioning with this kairomone via a reward. Finally, experiments on dispersal and edge behavior revealed that T. dubius and one of its common prey, the bark beetle I. grandicollis, exhibit a different pattern of movement within red pine stands and surrounding habitats. First, adult T. dubius can disperse substantially further (50% dispersed beyond 1.54 km) than I. grandicollis adults (50% dispersed beyond 0.13 km). Second, T. dubius was highly affected by the presence of an edge, showing a step-like pattern with very predators caught in unsuitable areas. Conversely, the behavior of I. grandicollis was less limited by such a boundary. As with T. dubius, the root colonizer D. valens showed a strong edge behavior that was characterized by avoiding matrix zones.

The Eurasian spruce bark beetle Dendroctonus micans is a major pest of spruce which is expanding its range in France, Turkey, England and Wales. Its monospecific predator Rhizophagus grandis has followed naturally the bark beetle into most areas and, since the 1960s, has also been mass-produced and successfully released within newly infested locations.

In this scope, the development of an effective trapping method would be very useful to assess the bark-beetle presence at previously uninfested sites, or predator establishment after release or natural spread. We demonstrated the efficiency of oxygenated monoterpenes-baited kairomone traps to monitor R. grandis in various epidemiological conditions, including areas localized behind or at the limit of the pest’s distribution, or in areas where artificial releases were performed. Because the predator is strictly species-specific, another exciting possibility offered by the kairomone trapping is the indirect monitoring of the pest itself in areas of unknown status (e.g. areas under colonization, or considered as at risk at medium- term).

R. grandis is also considered as one of the most valuable natural enemies to strike aggressive North-American Dendroctonus species. In this respect, R. grandis has been recently applied in a neo-classical biological program against the red turpentine beetle D. valens, which invaded China from North America in the late 1990’s. In laboratory tests conducted on pine logs in the laboratory, or on living pine trees in the field, we demonstrated that R. grandis adults can successfully enter and reproduce into D. valens galleries.

In Europe, R. grandis is the only species regularly found in the brood systems of D. micans, where adults and larvae attack the gregarious larvae of their prey. In such enclosed systems, R. grandis’ functional response is therefore influenced by various interrelated components, such as the prey density, the predator density, or the prey distribution. Measuring the predator’s success in terms of larval survival and growth rates, we demonstrated the time spent by R. grandis larvae to wound and kill their prey to be the main factor limiting their development. This factor may be considerably influenced by the proportions of diseased, wounded or molting prey rise in the brood system, for instance as a result of an increase in prey density, or due to the presence of conspecific adults (which wound their prey but do not consume them entirely). Furthermore, our tests suggest that no cannibalism or noticeable intraspecific competition occurred between R. grandis larvae, whereas some lighter mode of competition probably took place.

R. grandis also exhibits a reproductive numerical response to prey density, which mainly relies on the perception of chemical stimuli and inhibitors released in the bark beetle brood system. In the current study, we developed a non-destructive approach to follow the dynamics of volatile compound production, using sequential sample collection on SPME fibers. Our tests demonstrated that the larval activity of D. micans or D. valens strongly influences the release of some oxygenated monoterpenes. However, our attempts to correlate the relative quantities of some identified chemicals to offspring production were less successful as it concerns the identification of potential oviposition stimuli and inhibitors.

The problematic rose by the progression of D. micans, as well as detailed results of each of the described above studies are discussed in the two published papers and the three manuscripts forming this thesis. Bringing all these studies together, several perspectives are also presented in the general discussion.

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Ravageur des épicéas, Dendroctonus micans est toujours en voie d’extension en France, en Turquie, en Angleterre et au Pays de Galles. Dans la plupart de ces zones, le dendroctone est accompagné de manière naturelle par son prédateur monospécifique, Rhizophagus grandis. Depuis les années 1960, le prédateur a également fait l’objet d’une production de masse et de programmes de lâchers dans les zones d’arrivée récente du scolyte.

Dans le cadre de la lutte biologique contre D. micans, les gestionnaires forestiers doivent donc estimer au plus tôt la présence du ravageur dans des zones jusque là indemnes, mais également vérifier l’établissement du prédateur par progression naturelle ou résultant d’introductions délibérées. Dans la présente étude, nous avons démontré l’efficacité de pièges d’interception appatés à l’aide de monoterpènes oxygénés pour la capture de R. grandis. Celle-ci s’est faite dans différentes conditions épidémiologiques, incluant notamment des zones situées en arrière du front de progression du scolyte et des zones où des lâchers artificiels ont été réalisés. Comme R. grandis est strictement inféodé au dendroctone, un autre avantage de la technique est la possibilité de réaliser un dépistage indirect du ravageur dans les zones où son statut est incertain (zones en cours de colonisation, ou considérées comme à risque à moyen terme).

Par ailleurs, R. grandis est également considéré comme un des meilleurs ennemis naturels potentiels pour lutter contre d’autres espèces de Dendroctonus aggressifs. Dans cette optique, R. grandis a été récemment utilisé dans un programme de lute biologique contre D. valens, ravageur invasif arrivé en Chine dans la fin des années 1990 en provenance d’Amérique du Nord. Nous avons démontré la capacité de R. grandis à s’introduire et à se reproduire dans les galeries de D. valens lors de tests de laboratoire, mais aussi sur des arbres vivants en pinèdes.

En Europe, R. grandis est strictement inféodé aux galeries de D. micans, où larves et adultes du prédateur s’attaquent aux larves grégaires du scolyte. Dans ce système clos, la réponse fonctionelle de R. grandis est influencée par plusieurs facteurs étroitement corrélés, la densité de proies, la densité de prédateurs, et la distribution des proies. En mesurant l’efficacité de prédation de R. grandis en termes de survie des larves et de taux de croissance, nous avons démontré l’influence sur leur développement du temps passé par les larves à blesser et à tuer leurs proies. Ce facteur est par ailleurs fortement dépendant de la proportion de larves malades, blessées ou en cours de mue au sein du système ;une proportion qui peut augmenter en réponse à une augmentation de la densité de proies, ou lorsque des adultes sont présents (ceux-ci blessent les proies mais ne les consomment pas entièrement). Enfin, nos tests suggèrent qu’il n’existe que peu de cannibalisme ou de compétition intraspécifique entre larves de R. grandis, tandis que des modes de compétition moins importants prennent vraisemblablement place.

R. grandis présente également une réponse numérique reproductive à la densité de proies disponibles, principalement basée sur la perception de stimuli et d’inhibiteurs présents dans les galeries du scolyte. Par la collecte de composés volatils présents dans ces systèmes à l’aide de fibres SPME, nous avons développé une méthode non-destructive pour suivre la dynamique de production des médiateurs chimiques. Nos tests ont démontré que l’activité des larves de D. micans ou D. valens influence fortement la dynamique de production de certains monoterpènes oxygénés. En revanche, il n’a pas été été possible de corréler les différents composés identifiés au nombre de larves de R. grandis présentes dans le système.

La problématique soulevée par la progression de D. micans, de même que les résultats détaillés de chacune des études décrites ci-dessus sont discutés dans les deux papiers publiés et les trois manuscrits formant cette thèse. Les différentes perspectives apportées par ce travail sont également présentées dans la discussion générale.

Doctorat en Sciences agronomiques et ingénierie biologique
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Approximately 40% of Japanese forests are softwood plantations consisting of trees such as Japanese cedar (Cryptomeria japonica), Japanese cypress (Chamaecyparis obtusa), and several varieties of pine (Pinus spp.). Policies and programs related to forest pests and diseases are important for growing forest plantations. Damage caused by the pine bark beetle (Monochamus alternatus) has been a long-standing problem in Japan. Forest damage caused by the pine bark beetle was first found in Nagasaki Prefecture in 1905. Since then, the area of damage has expanded gradually to all prefectures. Damage caused by pine bark beetles became serious during and just after the end of the Second World War. In 1950, the Natural Resource Section of the General Headquarters of the Allied Forces, Supreme Commander for the Allied Powers (GHQ/SCAP) made recommendations for how to control forest pests and diseases. The first act was enacted in 1950, although the control of forest pests was initially addressed as part of the first Forest Act of 1897. Several important reasons for why the Japanese government has failed to stop the expansion of the damaged area can be found in GHQ recommendations: the lack of coordinated programs, underutilization of damaged trees, and shortcomings of forest-management plans.

The southern pine beetle, Dendroctonus frontalis Zimmermann (Coleoptera: Scolytidae), is an economically important pest of pine forests in the southern United States (Price et al. 1992). This native bark beetle is able to attack and kill living trees, typically loblolly (Pinus taeda L.) or shortleaf (Pinus echinata Mill.) pine, through a process of mass attack coordinated by pheromones emitted by the beetle (Payne 1980). During the attack process, thousands of beetles bore through the outer bark of the tree and begin constructing galleries in the phloem layer. Trees can respond to beetle attack by exuding resin from a network of ducts, but the large number of simultaneous attacks usually overcomes this defense, literally draining the resin from the tree. Oviposition and brood development then occur in the girdled (and ultimately dead) tree. Once a tree is fully colonized the attack process shifts to adjacent trees, often resulting in a cluster of freshly attacked trees, trees containing developing brood, and dead and vacated trees (Coulson 1980). These infestations can range in size from a single tree to tens of thousands, although the latter only occur in areas where no control methods are applied. Approximately six generations can be completed in a year in the southern United States (Ungerer et al. 1999). Like many other forest insect pests, D. frontalis populations are characterized by a considerable degree of fluctuation. The longest time series available are Texas Forest Service records of infestations in southeast Texas since 1958 (figure 5.la). These data suggest that the fluctuations have at least some periodic component, with major outbreaks occurring at intervals of 7-9 years (1968, 1976, 1985, and 1992). A variety of different analyses, including standard time series analysis and response surface methodology (Turchin 1990, Turchin and Taylor 1992), suggest that D.frontalis dynamics are indeed cyclic and appear governed by some kind of delayed negative feedback acting on population growth (see chapter 1). This effect can be seen by plotting the realized per-capita rate of growth (R-values) over a year against population density in the previous year (figure 5.1b).

Historically the boreal forest has experienced major changes, and it remains a highly dynamic biome today. During cold phases of Quaternary climate cycles, forests were virtually absent from Alaska, and since the postglacial re-establishment of forests ca 13,000 years ago, there have been periods of both relative stability and rapid change (Chapter 5). Today, the Alaskan boreal forest appears to be on the brink of further significant change in composition and function triggered by recent changes that include climatic warming (Chapter 4). In this chapter, we summarize the major conclusions from earlier chapters as a basis for anticipating future trends. Alaska warmed rapidly at the end of the last glacial period, ca 15,000–13,000 years ago. Broadly speaking, climate was warmest and driest in the late glacial and early Holocene; subsequently, moisture increased, and the climate gradually cooled. These changes were associated with shifts in vegetation dominance from deciduous woodland and shrubland to white spruce and then to black spruce. The establishment of stands of fire-prone black spruce over large areas of the boreal forest 5000–6000 years ago is linked to an apparent increase in fire frequency, despite the climatic trend to cooler and moister conditions. This suggests that long-term features of the Holocene fire regime are more strongly driven by vegetation characteristics than directly by climate (Chapter 5). White spruce forests show decreased growth in response to recent warming, because warming-induced drought stress is more limiting to growth than is temperature per se (Chapters 5, 11). If these environmental controls persist, projections suggest that continued climate warming will lead to zero net annual growth and perhaps the movement of white spruce to cooler upland forest sites before the end of the twenty-first century. At the southern limit of the Alaskan boreal forest, spruce bark beetle outbreaks have decimated extensive areas of spruce forest, because warmer temperatures have reduced tree resistance to bark beetles and shortened the life cycle of the beetle from two years to one, shifting the tree-beetle interaction in favor of the insect (Chapter 9).

The robin was twitching, tremoring, convulsing uncontrollably, and peeping occasionally. The student handed the bird to me, and in a few minutes it was dead in my hands. It was April 23, 1963, and I was in my laboratory at Dartmouth College in Hanover, New Hampshire, when the student walked in with the bird. A week earlier the elm trees of Hanover had been sprayed with the insecticide DDT to control the spread of Dutch elm disease by elm bark beetles. In the following weeks 151 dead birds filled my freezer, many of them exhibiting before they died the tremors that we later learned were typical of DDT poisoning. Four of us were conducting a small-scale study of the effects, if any, of the DDT spray program in Hanover. We were shocked by what was happening to the local birds, but we would have expected this reaction to DDT if we had read the scientific literature on earlier DDT spray programs on elm trees. We had not. We soon realized that we had rediscovered what other ornithologists had already reported from DDT spray programs in the American Midwest. We also soon learned that DDT was ineffective in preventing the spread of Dutch elm disease and that another procedure, sanitation without insecticides, effectively protected the elms. This DDT spray procedure was all costs and no benefits. Hundreds of towns were killing thousands or millions of birds while not protecting their elms. The whole thing struck me as absurd and tragic. It became a life-changing event for me. I decided that DDT was a chemical that had to be stopped, although I hadn’t the slightest idea where such a conclusion was going to lead. I was 33 years old and had become what in those days was usually called a conservationist. Now such people have been renamed “environmentalists.” I had a dubious beginning as such a person. When I was about seven and living in a northern suburb of Philadelphia, I came across a couple of snakes.

Numerical simulations were conducted to investigate the feasibility of predicting near field concentrations of a tracer gas within and above forest canopies. The current research is geared towards providing forest managers with a tool for developing anti-aggregation techniques to control the bark beetle. Several field experiments have been conducted in different forest canopies linking tracer gas concentration fields with meteorological and canopy parameters. Field experiment results are site and situation specific. Numerical simulations are far less expensive and allow for variation in virtually all flow parameters such as atmospheric stability, wind speed and direction, and turbulence intensity. As a first step, a CFD simulation has been used to study dispersion in a generic lodgepole pine forest canopy based on leaf area index (LAI) and stem density. Steady Reynolds Averaged Navier Stokes (RANS) solutions were computed using the k-ε and Reynolds Stress Model (RSM) turbulence closure models. These solutions provide insight into in-canopy dispersion; however they do not fully capture the dynamics of the flow. The current work uses large eddy simulation (LES). LES resolves large flow dominated eddies while modeling smaller eddies using a sub grid scale model. Unsteady LES, can be used to capture the dynamics of flow within a canopy, including large rolling eddies above the canopy, bursting and sweeping within the canopy, multiple shear layers, and drainage flows.