针叶树小蠹对非寄主挥发物的嗅觉识别和行为反应研究评述

针叶树小蠹在扬飞扩散过程中,要有效地依靠嗅觉机制鉴别寄主和非寄主植物的气味,以便在行为反应方面快速定位寄主,降低不利因素对种群的影响。不同的非寄主植物挥发物可以从生境、树种及个体水平上发挥不同的驱避作用。综述非寄主挥发物的范畴和作用、小蠹虫的嗅觉识别及行为反应,提出加强国内针叶树小蠹非寄主挥发物研究与开发的方向,以期为小蠹虫综合管理技术开发提供借鉴和参考。     

小蠹虫信息化合物作用研究概述

小蠹虫具有复杂的种间和种内信息化合物,其主要作用是调节和控制对寄主树木的选择、入侵及危害,进而调控小蠹虫在森林生态系统内的种群密度和生态分布。本文简要概述了信息化合物在小蠹虫寄主选择、入侵、定居、繁殖等一系列行为中的作用,并进一步阐述了小蠹虫寄主选择理论,重点阐明了寄主与非寄主植物挥发物的作用。     

小蠹科害虫化学信息物质及其对侵害寄主等行为的影响

在综合大量国内外研究成果的基础上,就松树寄主气味、非寄主气味、小蠹虫信息素以及其他化学信息物质对调控小蠹科害虫侵袭寄主林木等的行为,作了系统详细的论述。     

小蠹虫伴生菌研究概况

小蠹虫伴生菌是在植物-小蠹虫-伴生菌系统中,削弱树木抗性、协助小蠹虫侵害的重要因子。伴生菌与小蠹虫的联系和相互适应是在长期协同进化中形成的。寄主树木对伴生菌的抗御作用主要是通过物理防御、化学防御进行的。而伴生菌对寄主树的侵害又常常是与小蠹虫协同作用的。因此,揭示伴生菌在小蠹虫相互生态联系,探讨伴生菌在植物-小蠹虫-伴生菌系统中所扮演的角色,对于防治小蠹虫危害、保护寄主树木具有重要意义。     

应用信息化合物防治小蠹虫的研究进展

小蠹虫是世界上重要的森林害虫,常造成森林大面积死亡.小蠹虫多存在于高中山区且其生活隐蔽,利用传统的防治方法很难达到较好的效果.近年来利用化学信息物质防治小蠹虫成为国内外研究的热点.本文简要概述了化学信息物质,特别是寄主次生性物质和小蠹虫信息素对小蠹虫寄主选择行为的影响,以及国内外应用化学信息物质防治小蠹虫的进展.     

云杉八齿小蠹化学生态的研究进展(鞘翅目,小蠹科)

本文对云杉八齿小蠹化学生态的研究进展进行了综述。云杉八齿小蠹在中欧的大面积爆发激发了广大林业科学工作者对其化学生态学的广泛研究。在寻找寄主的过程中,雄性找到寄主后释放化学信息物质吸引雄性和雌性的聚集。开始发起进攻的小蠹在找寻寄主的过程中能利用寄主树脂作为利己素,并能够在体内解毒后合成可被利用的化学信息物质。在云杉八齿小蠹雄性的中肠中,我们发现9种单萜类化合物,主要是反式马鞭草烯醇和2-甲基3-己烯2-醇,这两种物质被认为是云杉八齿小蠹聚集信息素的主要成分。小蠹二烯醇的含量虽然很低但是能显著提高马鞭草烯醇和2-甲基3-己烯2-醇的野外诱集作用。马鞭草烯酮和小蠹烯醇是反聚集信息素,在调节攻击密度和树皮下的种群密度的过程中起着重要的作用。本文主要综述了寄主抗性、信息素成分和小蠹虫行为之间的关系。野外生测的结果证明利用信息素防治云杉八齿小蠹是行之有效的防治方法。     

牛背梁自然保护区植物群落和针叶树小蠹虫群落物种多样性研究

对牛背梁自然保护区的山麓农田-侧柏群落(群落Ⅰ),油松-华山松-锐齿栎群落(群落Ⅱ),油松-华山松-栓皮栎群落(群落Ⅲ),油松-华山松-辽东栎群落(群落Ⅳ),巴山冷杉-落叶松群落(群落Ⅴ),红桦-牛皮桦-巴山冷杉群落(群落Ⅵ)6种类型群落的高等植物及其针叶树小蠹虫的多样性等进行研究.结果表明:在山麓灌丛群落至针阔混交林群落之间,物种多样性指数、群落均匀度指数和物种丰富度呈递增趋势;在中山针阔混交林群落与巴山冷杉和落叶松群落、红桦-牛皮桦-巴山冷杉群落之间,物种多样性指数、群落均匀度指数和物种丰富度呈下降趋势.从总体来看,森林植被的多样性程度高于灌丛和草甸,阔叶林则高于针叶林.群落优势度指数的变化规律则与之相反.不同森林群落类型中针叶树小蠹虫的多样性、均匀度和优势度研究结果表明:针叶树小蠹虫群落物种多样性指数和均匀度由高到低的排列顺序依次为:群落Ⅱ、Ⅲ、Ⅳ、Ⅴ、Ⅵ、Ⅰ;针叶树小蠹虫群落优势度的变化趋势与之相反.不同类型的森林植物群落中,针叶树小蠹虫群落结构及多样性存在明显差异.生态环境复杂,植被丰富的中山针阔混交林群落中,针叶树小蠹虫多样性和均匀度指数均较大,但优势度指数低,小蠹虫大发生的几率较小.加强管理,促进森林生态系统的多样性和稳定性,对提高植物群落自身调控害虫种群数量具有重要意义.     

树木抗性与小蠹虫生存策略的进化

小蠹虫对寄主树木的入侵危害不仅受到树木原生性抗性的直接影响 ,而且小蠹虫在寄主树木组织内的定居繁衍也受到寄主树木诱导性抗性的制约。寄主树木抗性的强弱决定小蠹虫入侵寄主树木的数量 ,也促使小蠹虫与特定的真菌共生 ,真菌通过对寄主树木生理生化代谢的影响 ,以削弱寄主树木抗虫性 ,从而为小蠹虫在寄主树木组织内的定居创造必备的生存条件。同时小蠹虫为适应真菌的共生和寄主树木的抗虫性 ,促使小蠹虫生存繁衍策略高度进化 ,从而形成小蠹虫—共生真菌—寄主树木三者协同进化共生的关系     

小蠹虫伴生长喙壳类真菌研究进展

小蠹虫与多种微生物具有密切的关系,其中小蠹虫与伴生真菌种间关系存在着相对稳定性和依赖性.在小蠹虫蛀干过程中,伴生真菌随蠹虫携带进入树木木质部和韧皮部,通过消耗寄主的养料,堵塞木质部筛管,破坏韧皮细胞,对寄主树木产生危害作用.在小蠹虫众多的伴生真菌里,尤以长喙壳类真菌(ophiostomatoid fungi)在小蠹虫种群建立、协同克服寄主抗性、引起树木病害等方面起到重要作用,带来直接和间接的经济损失.该文对小蠹虫与长喙壳类真菌伴生关系、长喙壳类真菌种类等做一综述.     

秦岭华山松小蠹生态位研究

通过对秦岭林区海拔１６００～２２００ｍ的中山地带华山松小蠹虫种类和生态位的研究,结果表明：在秦岭林区入侵危害寄主华山松的小蠹虫有１９种,其中能构成竞争和共存的小蠹虫主要有１１种。虽然,这１１种小蠹虫具有各自不同的生态位宽度,且存在不同程度的生态位重叠,但各小蠹虫可依据其对寄主树木营养和空间需求的不同、种群密度的相互制约,以及入侵寄主树木时序的差异达到竞争的平衡和共存。秦岭华山松小蠹生态系统的建立,首先是由华山松大小蠹入侵健康华山松,并通过携带蓝变真菌入侵寄主树木,迅速克服寄主树木抗性系统,使寄主树木树势衰弱;其次松六齿小蠹、暗额星坑小蠹和松十二齿小蠹,作为秦岭华山松的主要次期性小蠹,迅速入侵衰弱的寄主华山松;最后其它次期性小蠹虫入侵寄主树木,利用寄主华山松剩余营养和空间。从而实现秦岭华山松立木小蠹生态系统的动态稳定。     

A review： chemical ecology of Ips typographus （Coleoptera, Scolytidae）

Chemical ecology of the spruce bark beetle lps typographus （L.） was reviewed. The outbreak of 1.typographus in central Europe triggered extensive research on chemical ecology, Males initiate host location and produce semiochemicals which attract both males and females, A successful mass attack must first overcome the resistance of the host tree. Pioneer I. typographus evolved to use the resin flow of host trees as kairomones in host location, and synthesized semiochemicals initially to detoxify the resin. If small bark beetle populations infest healthy trees, mass attack is prevented by host resistance, Nine monoterpene alcohols were found in male hind-guts, including cis-verbenol （cV） and 2-methyl-3-buten-2-ol （MB） which are regarded as primary aggregation pheromones, and a low proportion of lpsdienol （ld） which increases attractiveness of cV and MB, Verbenone （Vn） and Ipsenol （le） are anti-aggregation pheromones, that play important roles in adjusting attack density and insect density under the bark. Non-host volatiles are repellent to L typographus, so that beetles do not waste energy boring into non-host trees. The relationship between host resistance, pheromone compounds and behavior, non-host volatiles, bioassays and mass trapping are reviewed, Results of field bioassays stressed that traps baited with specific pheromones could be used as a reasonable protection measure.     

Test von Monoterpenen als Zusatz zu Pheroprax® bzw. Chalcoprax® in Pheromonfallen zum Fang des Buchdruckers,Ips typographus L. bzw. des Kupferstechers,Pityogenes chalcographus L. (Col., Scolytidae)

Experiments on monoterpenes in combination with Pheroprax^® and Chalcoprax^® in pheromone traps for catching the bark beetlesIps typographus L. andPityogenes chalcographus L. (Col., Scolytidae) It is highly evident that monoterpenes of the host tree serve as olfactory stimulants for “pioneer-beetles” colonizing first a tree before any bark-beetle-produced aggregation pheromones can lure those beetles to the host. The findings ofRedemann (1993) about a significant increase of spruce engraver catches by addition of both (?)-alpha-pinen and (+)-limonen simultaneously to pheroprax-baited bark beetle traps (Pheroprax^R being the aggregation pheromone ofIps typographus) induced us to conduct field experiments testing the same host volatiles in comparable bark beetle traps baited with either Pheroprax^® or Chalcoprax^® (aggregation pheromone ofPityogenes chalcographus). Despite we used methods considering the influence of place and time on the bark beetle flight activities, in 3 experiments with 30 repetitions in total no monoterpene-induced enhancement oftypographus-catches could be found; the same was true withP. chalcographus (2 experiments, 18 repetitions in total). With respect to the important role of host volatiles also from a practical point of view, it is strongly recommended to repeat experiments like these under different conditions to reveal the reasons of the different findings.     

Overview of development of an anti-attractant based technology for spruce protection against Ips typographus: From past failures to future success

We describe a series of experiments on the protection of Norway spruce (Picea abies (L.) Karst.) logs and standing trees against Ips typographus L. (Coleoptera: Scolytidae). The tested protective materials were applied 1) at point sources and 2) area-covering using zeolite-based dispensers. We performed 11 terrestrial and one aerial application (using a helicopter) between 1992 and 2001. We gradually replaced polyethylene-bag verbenone dispensers by composite dispensers releasing verbenone and non-host volatiles (NHV: green leaf volatiles, trans-conophthorin, and C8-alcohols). Many experiments did not give statistically significant results, due to the low number of true replicates and the spatial heterogeneity of beetle attacks. The protection of horizontal objects like logs or lying trees with anti-attractants was not always successful. In the future, positive results can be expected with an area covering spray applied by zeolite dispensers containing mixtures of verbenone and NHV. A similar approach could probably be used in aerial spraying of anti-attractants. We have succeeded in our attempt to protect zones of standing spruce trees from I. typographus attacks. We used dispensers with a combination of verbenone and a full blend of NHV compounds. The protection is not complete, but in both 2000 and 2001, the treatment significantly decreased the probability of the insect attack on trees by 60–80?%.     

Effects of volatiles of non-host plants and other chemicals on oviposition of <Emphasis Type="Italic">Monochamus alternatus</Emphasis> (Coleoptera: Cerambycidae)

The response of adult females of Monochamus alternatus to volatiles of non-host plants was evaluated using a Y-tube olfactometer. Eight non-host plant species were tested, namely Broussonetia papyrifera, Cedrus deodara, Firmiana simplex, Metasequoia glyptostroboides, Platycladus orientalis, Populus xiaohei, Salix babylonica and Sophora japonica. The volatiles of all test species, except those of C. deodara, repelled the females in the olfactometer bioassays. Oviposition by adult females on the host plant, Pinus massoniana, was deterred by the volatiles of non-host plants and by some other chemicals. Volatiles of P. orientalis had the strongest oviposition-deterring activity and those of S. japonica the least. Volatiles of C. deodara had no influence on oviposition of the females. Selected chemicals, namely benzaldehyde, citronellal, essential oil of Mentha spicata, eucalyptol, nerolidol, nicotine and salicylaldehyde, strongly deterred the females from oviposition, and nicotine caused maximum decrease in egg-laying and scar-excavation.     

Colonisation of native and exotic conifers by indigenous bark beetles (Coleoptera: Scolytinae) in France

Planting exotic conifers offers indigenous forest insects an opportunity to extend their host range and eventually to become significant pests. Knowing the ecological and evolutionary modalities driving the colonisation of exotic tree species by indigenous insects is thus of primary importance. We compared the bark beetle communities (Coleoptera: Curculionidae, Scolytinae) associated with both native and introduced conifers in France. The aim of our study was to estimate the influence of both host- and insect-related factors on the beetles’ likelihood to shift onto new hosts. We considered the influence of host origin (i.e. native vs. exotic), host tree species identity, tree bark thickness and tree taxonomic proximity, as well as insects’ host specificity. A field inventory using trap trees was carried out in two regions in France (Limousin and Jura) during two consecutive years (2006 and 2007) on three European native conifer species [Norway spruce (Picea abies); Scots pine (Pinus sylvestris) and European Silver-fir (Abies alba)] and five North American [Sitka spruce (Picea sitchensis); Eastern white pine (Pinus strobus); Grand fir (Abies grandis); Douglas fir (Pseudotsuga menziesii) and Western red cedar (Thuja plicata)]. A total of 18 indigenous and 2 exotic bark beetle species were collected. All exotic conifer species were colonised by indigenous bark beetle species and no significant difference was observed of the cumulated species richness of the latter between native and exotic tree species (13 vs. 14, P < 0.05). The ability of indigenous bark beetles to shift onto exotic conifers appeared to strongly depend on host species (significantly structuring bark beetle assemblages), the presence of phylogenetically related native conifer species and that of similar resources, in combination with insect host specificity. Host tree species status (native or exotic) also seemed to be involved, but its effect did not seem as essential as that of the previous factors. These findings are discussed in terms of adaptation, plasticity and practical aspects of forest management.     

Response of bark beetles and their natural enemies to fire and fire surrogate treatments in mixed-conifer forests in western Montana

Four treatments (control, burn-only, thin-only, and thin-and-burn) were evaluated for their effects on bark beetle-caused mortality in both the short-term (one to four years) and the long-term (seven years) in mixed-conifer forests in western Montana, USA. In addition to assessing bark beetle responses to these treatments, we also measured natural enemy landing rates and resin flow of ponderosa pine (Pinus ponderosa) the season fire treatments were implemented. All bark beetles were present at low population levels (non-outbreak) for the duration of the study. Post-treatment mortality of trees due to bark beetles was lowest in the thin-only and control units and highest in the units receiving burns. Three tree-killing bark beetle species responded positively to fire treatments: Douglas-fir beetle (Dendroctonus pseudotsugae), pine engraver (Ips pini), and western pine beetle (Dendroctonus brevicomis). Red turpentine beetle (Dendroctonus valens) responded positively to fire treatments, but never caused mortality. Three fire damage variables tested (height of crown scorch, percent circumference of the tree bole scorched, or degree of ground char) were significant factors in predicting beetle attack on trees. Douglas-fir beetle and pine engraver responded rapidly to increased availability of resources (fire-damaged trees); however, successful attacks dropped rapidly once these resources were depleted. Movement to green trees by pine engraver was not observed in plots receiving fire treatments, or in thinned plots where slash supported substantial reproduction by this beetle. The fourth tree-killing beetle present at the site, the mountain pine beetle, did not exhibit responses to any treatment. Natural enemies generally arrived at trees the same time as host bark beetles. However, the landing rates of only one, Medetera spp., was affected by treatment. This predator responded positively to thinning treatments. This insect was present in very high numbers indicating a regulatory effect on beetles, at least in the short-term, in thinned stands. Resin flow decreased from June to August. However, resin flow was significantly higher in trees in August than in June in fire treatments. Increased flow in burned trees later in the season did not affect beetle attack success. Overall, responses by beetles to treatments were short-term and limited to fire-damaged trees. Expansions into green trees did not occur. This lack of spread was likely due to a combination of high tree vigor in residual stands and low background populations of bark beetles.     

Zur Kenntnis desOrthotomicus tridentatus Egg. (Zedernborkenkäfer) einem Schädling derZedernwälder der Türkei

Summary O. tridentatus was an almost unknown bark beetle until this study. The bionomics, ecology, distribution and economic importance of this species are described here; and preventive and control measures are recommended. It seems to be confused withO. erosus because of its very close morphological similarity.O. tridentatus is a cedar bark beetle, but cedar ist not a host ofO. erosus. The former is distinguished from the latter by the acute angle between the 2nd and 3rd teeth at the end of its abdomen, by the slight difference in the pattern of its antennal club, and by the form of its brood burrows.O. tridentatus is the most dangerous bark beetle of the cedar forest. It is monophagous and is 2.8 to 3.6 mm long. Its burrows are generally S-shaped transverse or diagonal burrows with 2 egg galleries. If there is a large concentration of beetles in a small area of a tree trunk, the burrows can be longitudinal. The entire burrow ranges 3.6 to 15 cm (average: 7 cm) in length and 1.2 to 2.2mm in width and has 40 to 164 larval burrows, the average being about 90 burrows which are from 2 to 5,5 cm long. The distribution of this species corresponds closely to that of its host,Cedrus libanotica Link. However, during this study, the bark beetle was found only at altitudes ranging from 1300 to 1700 meters. It apparently has at least two generations each year, and hibernates as a young adult. The first flight of this species takes place in the later part of April.      

Types of bark beetle (Coleoptera: Scolytidae) infestation in spruce forest stands affected by air pollution, bark beetle outbreak and honey fungus (Armillaria mellea)

583 spruce stands in an area affected by air pollution and bark beetle outbreak in Eastern Slovakia were studied in 1996. According to bark beetle infestation of dominant and codominant trees, stands were classified into following types of spruce stand decline:Ips typographus-A,Ips typographus-B,Polygraphus poligraphus, I. typographus/P. poligraphus—A,I. typographus/P. poligraphus—B. The presence of attacked trees in forest edges, bark beetle spots and forest interior was the key important factor for the classification. Data from forest inventory and forest management evidence together with data on types of spruce stands decline were used in further analyses. Results shows that the distribution of forest stands classified into different types or uninfested stands is related mainly to host size and site quality. The percentage of spruce, exposition of stands and stand density showed significant effects. The mechanisms of spreading of studied bark beetle outbreak could be explained by direct effects of stress of trees caused by an abrupt increase of level of solar irradiation and by weakening of trees by the honey fungus.