赤松林上调力及群落结构对昆嵛山腮扁叶蜂的调控机制及作用

赤松作为昆嵛山腮扁叶蜂的主要寄主,赤松林上调力及群落结构对该食叶害虫种群密度的调控机制一直是昆嵛山区森林生态系统研究的核心问题之一,也是有害生物可持续控制的重要理论基础。本文把昆嵛山赤松林生态系统作为研究对象,以赤松林林分特征和以寄主植物的多样性、树种组成、邻近木和物种联结性为主的群落结构为主线,阐明了赤松林上调力及群落结构调控昆嵛山腮扁叶蜂的作用和机制,为研究昆嵛山腮扁叶蜂与寄主植物及其群落结构的关系开辟新思路和新领域,进而为真正实现森林食叶害虫的可持续控制提供依据。     

赤松纯林林分特征对昆嵛山鳃扁叶蜂发生量的影响

为探讨昆嵛山鳃扁叶蜂的发生与天然赤松纯林的林分特征及其自然分布的关系,依托昆嵛山森林生态定位研究站所设立的40块永久性样地,比较了群落自然演替13 a的赤松种群特征以及林内灌草多样性指数,并分析了虫口密度与赤松林分特征的关系。结果表明,13 a自然生长,赤松种群的径级分布和高度结构出现显著变化。赤松种群中占总数60%的林木径级为5 cmDBH≤25 cm,比1996年高51%;而约占67%的赤松个体高度在2 m到10 m之间,比1996年高57%。同时种群密度从1996年的平均超过13000株/hm2降到了2008年的平均2377株/hm2。赤松林内灌草Shannon多样性指数(H)和均匀度指数(JS)分别为2.50和0.79,分别低于1996年的(H)2.69和(JS)0.85,暗示赤松纯林生物多样性有降低的趋势。昆嵛山鳃扁叶蜂的虫口密度与赤松林分的郁闭度和赤松分布的海拔高度呈极显著相关关系、相关系数分别为R=0.931、P=0.002和R=0.924、P=0.003;与林分密度(R=0.780,P=0.038)、林木胸径(R=0.816,P=0.025)呈显著相关关系,而与树高以及树龄关系不显著。昆嵛山鳃扁叶蜂虫口密度的分布格局似乎非常符合"资源集中"假说。     

昆嵛山腮扁叶蜂发生与赤松纯林林分因子的关系

为了解林分因子与昆嵛山腮扁叶蜂发生的关系,为昆嵛山赤松纯林的营林措施和昆嵛山腮扁叶蜂的防控技术提供理论指导,本研究在昆嵛山区域选取林龄相对一致(34±2 a)的赤松纯林为对象,调查林分密度、郁闭度、草本盖度、树高、枝下高、胸径、冠幅7个林分因子以及昆嵛山腮扁叶蜂虫巢数。依据虫情指数的分布范围,将样地分为4组,通过方差分析和多重比较研究了4组样地中各项林分因子的差异性。采用逐步回归法,筛选出影响昆嵛山腮扁叶蜂虫情指数的关键因子,在此基础上通过偏相关分析和单因素方差分析法进一步阐明关键因子与虫情指数之间的关系。结果表明,7个林分因子在4组样地中均存在极显著的组间差异。逐步回归筛选出冠幅、枝下高和林分密度为影响昆嵛山腮扁叶蜂发生的关键因子,根据关键因子建立了虫情指数的线性预测方程。冠幅和枝下高对昆嵛山腮扁叶蜂虫情指数的影响均达到极显著正相关水平(P0.01),林分密度对昆嵛山腮扁叶蜂虫情指数的影响为极显著负相关(P0.01)。影响虫情指数的关键因子可能会因林分的不同而不一致,因此要根据关键因子调整林分结构,以达到控制虫害流行的目的。     

昆嵛山腮扁叶蜂与林下植物的物种联结性

为明确昆嵛山腮扁叶蜂(Cephalcia kunyushanica)的出现与林下植物是否关联,本研究在昆嵛山林区设置的50块永久样地内,应用样圆法进行了灌草层植物调查,并基于2×2联表,应用总体方差分析、χ2统计检验、Ochiai关联度指数、Dice指数、联结系数、共同出现百分率、点相关系数以及Cramer指数等指标,分别对昆嵛山腮扁叶蜂与邻近环境中所有195种林下植物种群的总体相关性、该昆虫与各植物构成的物种对之间联结显著性以及联结强度进行了测度。结果表明:(1)昆嵛山腮扁叶蜂与195个植物物种种群总体存在极显著的正相关;(2)该食叶昆虫与83个物种分别构成的物种对呈正联结关系,与另外112个物种分别构成的物种对呈负联结关系,负联结的比例高于正联结;(3)该昆虫与羊胡子草(Carex rigescens)等11种植物具有极显著的正联结关系,与宽叶苔草(C.siderosticta)等16种植物具有极显著的负联结关系。     

基于生物多样性保护的兴安落叶松与白桦最佳混交比例——以阿尔山林区为例

选取内蒙古阿尔山林区7种不同混交比例的松桦混交林,包括落叶松纯林、白桦纯林及5种不同比例的混交林为研究对象。对植物和昆虫种群丰富度、多样性进行了研究,并对种群在乔木层混交状况梯度下的分布进行了分析。结果表明:植物和昆虫种群分布受乔木层混交状况影响显著。同时,郁闭度能改变其分布状况及决定于松桦比例。兴安落叶松和白桦混交比例介于5∶5和7∶3之间时最有利于森林多样性,是进行林分疏伐的理想模式。     

应用物种指示值法解析昆嵛山植物群落类型和植物多样性

生物指示目前被广泛用于各类生态系统恢复的监测和评估、生态系统管理的政策与法规的制定等。本文的目的是采用物种指示值法初步解析昆嵛山植物群落类型及其植物多样性的特点,为进一步开展昆嵛山森林自然恢复评价和森林管理提供参考依据。物种累计曲线分析表明,从昆嵛山自然保护区40块永久性标准样地实际采到的物种占全部物种数（估计值ACE为131.26）的92.9%,显示抽样充分。采用主坐标分析将昆嵛山森林群落进一步划分为六种林分类型,通过物种指示值分析得到了各林分类型的指示物种。其中,黑松、山槐、麻栎是黑松林的指示物种;郁李、赤松、山胡椒是赤松林的指示物种;日本落叶松/刺杉林的显著指示物种包括日本落叶松、水榆花楸、刺杉、白檀和水蜡。针叶树－麻栎混交林的指示物种是麻栎;而针叶树－杂木混交林的指示物种构成复杂,如粘鱼须、楸树、华山松,等。阔叶林的物种组成很不相同,分别有枫香林、麻栎林、刺槐林、和水榆花楸林。综合比较不同林分类型乔灌草的整体多样性特点表明,6种林分类型的物种多度和物种丰富度无显著差异;阔叶林的Fisher α指数（P< 0.001）、Shannon–Wiener指数（P = 0.001）,Simpson指数（P = 0.034）与其它5种林分类型相比差异显著;其他五种林分类型的Fisher α指数和Shannon–Wiener指数无显著差异,但Simpson指数在这些林分间差异显著。昆嵛山林分类型的多样性与30年前树种引进和人工造林等人为干扰密不可分。     

黄土高原柴松群落空间结构

利用角尺度、大小比数和混交度3个林分空间结构参数,分析了黄土高原柴松(Pinus tabulaeformisf.shekannesis)群落的空间结构特征。结果表明:1)该林分的树种组成为柴松:辽东栎(Quercus liaotungensis)∶其他树种=7∶2∶1,林分密度为3064株.hm-2,柴松种群优势度明显。2)柴松林林分平均混交度为0.33,处于弱度混交到中度混交过渡时期,种间隔离程度较小。3)柴松林林分平均角尺度为0.57,整个林分呈团状分布。4)林分平均胸径大小比为0.52,平均树高大小比为0.53,林木大小比数的分布较为均匀。胸径大小比和树高大小比在反映林木生长优势方面具有一致性。     

树种相互作用、林分密度和树木大小对华北落叶松生产力的影响

本研究以塞罕坝地区华北落叶松纯林、白桦纯林和华北落叶松与白桦混交林为对象,每种林分类型设置2个林分密度(200～340和880～1100株·hm^-2)。基于树木大小分层取样,采集树芯样本668条。运用线性混合模型,分析了树种相互作用、林分密度和树木大小对华北落叶松生产力的影响。结果表明: 华北落叶松断面积生长量不同程度地受竞争、胸径、年龄和邻体密度的影响;在林分密度较高的混交林中,由于白桦对华北落叶松生长的促进作用,落叶松生产力发生明显的增益效应;在林分密度较低的混交林中,落叶松和白桦之间不发生相互作用,2个树种的生产力均低于相应的纯林;种内竞争是影响华北落叶松生产力的主要因素;树木大小对华北落叶松生产力产生积极的影响,但影响程度因林分密度和树种组成而异。适当增加林分密度和选择白桦作为混交树种可以提高华北落叶松生产力。     

林分结构对湖南栎类天然次生林林下植被生物量的影响

对湖南栎类天然次生林不同林分类型的林下植被生物量特征及其影响因素进行研究。结果表明: 将50块样地划分为甜槠-鹿角杜鹃混交林、亮叶水青冈-箭竹混交林、石栎-虎刺+山茶混交林、甜储+枹栎-鹿角杜鹃混交林、青冈栎-油茶+杜鹃混交林5种林分类型。5种林分的林下植被生物量均较低,不超过2.3 t·hm^-2。5种林分类型的林下灌木生物量没有显著差异,甜槠-鹿角杜鹃混交林草本层生物量显著低于其他4种林分类型。影响不同林分林下植被生物量的因素不同。在甜槠-鹿角杜鹃混交林,草本层生物量与郁闭度、角尺度呈显著负相关,林下植被总体生物量与开敞度呈显著正相关;在亮叶水青冈-箭竹混交林,灌木层生物量与林分的角尺度、郁闭度呈显著负相关,草本层生物量与林下植被总体生物量与混交度呈显著正相关;在石栎-虎刺+山茶混交林,草本层生物量与林分的聚集指数呈显著正相关;在甜储+枹栎-鹿角杜鹃混交林,灌木层生物量与林分的混交度呈显著负相关,草本层生物量与林分的株数密度呈显著正相关;而在青冈栎-油茶+杜鹃混交林,林分结构与林下生物量没有显著相关关系。对林下植被生物量进行调整,应该以调整林分水平分布格局为主,综合考虑林分遮盖程度与树种结构。     

塞罕坝自然保护区樟子松不同林分类型对昆虫群落多样性的影响

为研究近自然森林经营下的樟子松人工林不同林分类型对昆虫群落多样性的影响,本文以塞罕坝自然保护区内的樟子松纯林与混交林作为研究对象,对5种林分类型(樟子松、樟子松-落叶松、樟子松-白桦、樟子松-山荆子、樟子松-落叶松-白桦-山荆子-山刺玫)中的昆虫多样性进行分析.共获得昆虫标本9617头,隶属于7目70科195种,以双翅目、半翅目和鞘翅目为优势类群.5种不同林分类型中的昆虫物种均比较丰富,且各林分类型之间的昆虫群落结构均处于中等不相似水平,樟子松混交林内的昆虫物种数和个体数均高于纯林,但昆虫群落特征指数在不同林分类型之间无显著差异.主成分分析显示,植食性类群、捕食性类群和寄生性类群是构成昆虫群落结构的主要成分,且捕食性和寄生性等天敌类群对植食性类群的制约作用较强,各林分内昆虫群落结构均具有较高的稳定性.     

Habitat modification contributes to associational resistance between herbivores

Associational resistance, in which one species gains protection from its consumers by association with a competitor, is common among plants but has seldom been documented among insects. Here we show that parasitism of an aboveground herbivore, European corn borer, by its specialist parasitoid Macrocentrus grandii, is reduced 98% in the presence of a belowground herbivore, corn rootworm. We tested the hypothesis that this positive indirect interaction between corn rootworm and corn borer was mediated by corn rootworm’s influence on the habitat: plant height was diminished by 33% and plant density by 20%, resulting in a more open habitat. We found that M. grandii showed a two- to five-fold preference for dense versus open habitats, and that experimental reduction of plant density in the absence of corn rootworm reduced M. grandii parasitism of corn borer by 13%. This result supports the presence of a habitat modification effect as a contributing factor to associational resistance for corn borer. We argue that associational resistance may be more common among phytophagous insects than previously appreciated.     

Scale-dependent neighborhood effects: shared doom and associational refuge

A resource’s susceptibility to predation may be influenced by its own palatability and the palatability of its neighbors. We tested for effects of plant chemical defenses on seed survival by manipulating the frequency of palatable and less palatable sunflower seeds in food patches subject to harvest by fox squirrels (Sciurus niger) and gray squirrels (Sciurus carolinensis). We varied resource distributions at three scales: among stations (aggregates of patches ca. 50 m apart), among patches immediately adjacent to each other, and within patches. When food patches were segregated into high-palatability and low-palatability stations (Experiment 1), seeds suffered greater mortality at stations with high levels of palatable seeds. In the same experiment, within patches, squirrels selected strongly for palatable seeds over less palatable seeds. When high- and low-palatability food patches were placed together at the same stations (Experiment 2), increasing densities of co-occurring palatable seeds amplified the mortality of less palatable seeds, indicating “shared doom.” When palatable and less palatable seeds were partitioned into micropatches (Experiment 3), associational effects disappeared, as predicted. Furthermore, selectivity in less palatable patches increased as the initial densities of palatable seeds increased, and selectivity in palatable patches decreased as the initial densities of less palatable seeds increased. Foraging theory predicts associational effects among prey that vary in palatability. Our results show how the type and magnitude of associational effects emerge from the interplay among the spatial scale of prey heterogeneity, the diet selection strategy, and the scale-dependent foraging responses of the consumer.     

Effects of spatial distribution on plant associational defense against herbivory

Several studies have shown that consumption of a focal plant by herbivores depends not only on its own defense traits but also on the characteristics of the neighboring plants. A number of studies have reported on plant associational defense in relation to neighboring plant palatability but the effect of the spatial distribution of the focal plant within patches of different neighboring plants has received less attention. We conducted a manipulative experiment to determine whether and how spatial distribution of focal plants affects the associational defense between plant species. In our experimental setup sheep encountered two patches varying in spatial distribution of the focal plant within patches (dispersed or clumped) and patch quality, good patch and bad patch, where the focal plant, Lathyrus quinquenervius, was neighbored to high- (Chloris virgata) or low-palatable (Kalimeris integrifolia) species, respectively. Results showed that, when focal plants were dispersed within both patches, the risk of attack was significantly lower for focal plants in the patches with low- than high-palatable neighbors, indicating associational defense. Alternatively, when focal plants were clumped within both patches, they were consumed in bad-patch as much as in good-patch plots, which indicates the absence of associational defense. However, if the focal plants have different spatial distributions in the two patches (dispersed in good-patch and clumped in bad-patch or vice versa), sheep foraging success for focal plants was greatly reduced in dispersed spatial pattern irrespective of the palatability of neighboring plants. Therefore, we concluded that spatial distribution is as important as traits of neighboring plants in predicting vulnerability of the focal plant to grazing by generalist herbivores. The outcome of plant associational defense for different types of neighborhood strongly depends on the magnitude of herbivore foraging selectivity between and within patches, which further depended on the contrasts between plant species or between patches.     

Non-host volatiles mediate associational resistance to the pine processionary moth

An increasing body of evidence indicates that the association between different plant species may lead to a reduction in insect herbivory, i.e. associational resistance. This might be due to a top–down regulation of herbivores by increased numbers of natural enemies or to a disruptive bottom–up influence of lower host plant accessibility. In particular, the richer plant communities release more diverse plant odours that may disturb olfactory-guided host choice and mating behaviour of insect herbivores, i.e. the “semiochemical diversity hypothesis”. However, this hypothesis has been rarely tested experimentally in natural habitats, notably forest ecosystems. We tested the effects of non-host volatiles (NHV) on mate and host location by the pine processionary moth (PPM) at the scale of individual pine trees with branches of non-host tree (birch) at their base. Pheromone trap catches and the numbers of larval nests were both reduced by non-host presence under treated pine trees, confirming an associational resistance mediated by NHV. In both males and females, the antenna could detect several birch volatiles, including methyl salicylate (MeSa). MeSa inhibited the attraction of the PPM male to pheromone traps, as did bark and leaf chips from birch trees. Our test of three doses of MeSa at the habitat scale (50 m forest edges) showed that the reduction in the numbers of male PPM captured in traps and in larval nests was MeSa dose-dependent. These results show that odours released by deciduous non-host trees can reduce herbivory by a forest defoliator in conifers, providing support to the “semiochemical diversity hypothesis” as a mechanism of associational resistance.     

Mating affects resource selection and modulates associational effects between neighbouring resources

Associational effects occur when the attack rate on a resource depends on neighbouring resources in the environment. These effects are predicted to result from mismatches experienced by the consumer organism in resource selection along hierarchical search levels. As resource selection depends on sensory information used during search behaviour, we expected that different physiological states of an insect might modulate the outcome of associational effects due to differences in resource selection. We used Drosophila melanogaster, as a model organism for olfactory‐guided behaviour in insects, to study the effects of mating induced behavioural changes on associational effects between two alternative resources. We found that mating has no effect on the ability of D. melanogaster to locate resource patches, but rather affects the perception of the resources within the patch. Consequently, we only found associational effects in the experiments with unmated females and not in the experiments with mated females. Our results suggest that the lack of associational effects for mated females resulted from changes in the use of short‐range olfactory cues, leading to random selection among the resources. In conclusion, our results suggest that the physiological state of an insect modulates associational effects by affecting resource selection rates within the patch.     

Effects of associational resistance and host density on woodland insect herbivores

1. Specialist herbivores often become less abundant per unit of host tissue as host density increases (resource dilution). They usually become less abundant when non-host species are mixed with their host plants (associational resistance). Most studies of these trends have involved herbaceous host plants and have not examined both trends for the same herbivores. 2. Three hypotheses were tested for the response of insect specialists to host plant density: resource concentration, plant apparency and resource dilution. Two hypotheses were tested for the response of herbivores to non-host plants: associational resistance and plant apparency. 3. From 1992 to 2007, I examined the responses of three monophagous insect herbivores to the densities of their host, Pinus edulis, and of two non-hosts, Pinus ponderosa and Juniperus spp. 4. Herbivore loads increased with host density, though the correlations were weak and often variable between generations. These results were consistent with the resource concentration and plant apparency hypotheses, but not with resource dilution. 5. Herbivore loads decreased as non-host density increased, consistent with the associational resistance hypothesis. This and other studies have shown that associational resistance is important in many types of plant communities. 6. The absence of resource dilution on woodland trees contrasted with studies of herbaceous host plants. Further comparisons of woody and herbaceous host plants are needed to elucidate the reasons for this difference.     

Searching behavior and associational response ofZetzellia mali (Acarina: Stigmaeidae)

Experimental tests of the hypothesis that natural selection favors predators that search and select prey species that are more profitable are reported. The predator studies was the stigmaeid mite,Zetzellia mali (Ewing). It was observed thatAculus schlechtendali (Nalepa) andPanonychus ulmi (Koch) were more profitable prey thanTetranychus urticae Koch. However,Z. mali attacks prey independently of their nutritional value. Behavioral observations and experiments documented thatZ. mali does not detect kairomones of the prey species. Therefore, prey location is due only to random encounters. Apparently,Z. mali does not search and select prey that are more profitable. The correlation coefficients ofZ. mali, A. schlechtendali, andP. ulmi were investigated in an apple tree. Results revealed that the association depends on season and type of prey.Zetzellia mali andA. schlechtendali were positively associated in the spring (r=0.3,P<0.05) and fall (r=0.36,P<0.01).Zetzellia mali andP. ulmi were not statistically associated. The term associational response is proposed as an ecological process that quantifies the association of predator and prey. The associational-response data suggest thatZ. mali, in the natural ecosystem, responds more to the density ofA. schlechtendali than to the density ofP. ulmi.     

Testing associational resistance against pine weevils mediated by Lasius ants attending conifer seedlings

Natural enemies attracted to plants may provide those plants with protection against herbivores but may also protect neighbouring plants, that is through associational resistance. Ant attendance may be an important mechanism for the occurrence of such effects because ants can reduce the damage caused by a wide variety of herbivorous insects. Ants have been shown, in a previous field experiment, to decrease the damage caused by the pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), a pest species that causes high seedling mortality in forest regeneration areas. In this study, we specifically tested whether seedlings planted close to ant‐attended seedlings experience associational resistance. We did this under laboratory conditions using the ant species Lasius niger (L.) (Hymenoptera: Formicidae). The feeding damage by pine weevils was significantly reduced on seedlings attended by ants. The neighbouring seedlings, however, did not experience associational resistance. Nevertheless, some associational effects were observed as the number of weevils recorded on both ant‐attended and neighbouring seedlings was significantly lower compared with ant‐excluded seedlings.