Experimental evidence for apparent competition in a tropical forest food web

The herbivorous insects of tropical forests constitute some of the most diverse communities of living organisms. For this reason it has been difficult to discover the degree to which these communities are structured, and by what processes. Interspecific competition for resources does occur, but its contemporary importance is limited because most pairs of potentially competing insects feed on different host plants. An alternative way in which species can interact is through shared natural enemies, a process called apparent competition. Despite extensive theoretical discussion there are few field demonstrations of apparent competition, and none in hyper-diverse tropical communities. Here, we experimentally removed two species of herbivore from a community of leaf-mining insects in a tropical forest. We predicted that other species that share natural enemies with the two removed species would experience lower parasitism and have higher population densities in treatment compared with control sites. In both cases (on removal of a dipteran and a coleopteran leaf-miner species) we found significantly lower parasitism, and in one case (removal of the dipteran) we found significantly higher abundance a year after the manipulation. Our results suggest that apparent competition may be important in structuring tropical insect communities.     

Predator diversity dampens trophic cascades

Food web complexity is thought to weaken the strength of terrestrial trophic cascades in which strong impacts of natural enemies on herbivores cascade to influence primary production indirectly. Predator diversity can enhance food web complexity because predators may feed on each other and on shared prey. In such cases, theory suggests that the impact of predation on herbivores relaxes and cascading effects on basal resources are dampened. Despite this view, no empirical studies have explicitly investigated the role of predator diversity in mediating primary productivity in a natural terrestrial system. Here we compare, in a coastal marsh community, impacts of arthropod predators on herbivores and plant productivity between a simple food web with a single predator species and a complex food web with a diverse predator assemblage. We show that enhancing predator diversity dampens enemy effects on herbivores and weakens trophic cascades. Consequently, changes in diversity at higher trophic levels can significantly alter ecosystem function in natural systems.     

Ecosystem-level patterns of primary productivity and herbivory in terrestrial habitats

Ecosystems are structurally organized as food webs within which energy is transmitted between trophic levels and dissipated into the environment. Energy flow between two trophic levels is given by the amount of production at the lower level and by the proportion of production that is consumed, assimilated and respired at the higher level. Considerable evidence indicates that food-web structure varies predictably in different habitats, but much less is known about quantitative relationships among food web fluxes. Many of the energetic properties of herbivores in African game parks are associated with rainfall and, by inference, with net primary productivity. Respiratory costs per unit production at the consumer trophic level are higher for homeotherms than for heterotherms. Plant secondary chemicals affect herbivore dietary choices and the allocation of plant resources to those chemicals varies with resource availability. How these phenomena are translated into ecosystem fluxes is unknown. We present evidence that herbivore biomass, consumption and productivity are closely correlated with plant productivity, suggesting that the latter is a principal integrator and indicator of functional processes in food webs.     

Low host specificity of herbivorous insects in a tropical forest

Two decades of research have not established whether tropical insect herbivores are dominated by specialists or generalists. This impedes our understanding of species coexistence in diverse rainforest communities. Host specificity and species richness of tropical insects are also key parameters in mapping global patterns of biodiversity. Here we analyse data for over 900 herbivorous species feeding on 51 plant species in New Guinea and show that most herbivorous species feed on several closely related plant species. Because species-rich genera are dominant in tropical floras, monophagous herbivores are probably rare in tropical forests. Furthermore, even between phylogenetically distant hosts, herbivore communities typically shared a third of their species. These results do not support the classical view that the coexistence of herbivorous species in the tropics is a consequence of finely divided plant resources; non-equilibrium models of tropical diversity should instead be considered. Low host specificity of tropical herbivores reduces global estimates of arthropod diversity from 31 million (ref. 1) to 4 6 million species. This finding agrees with estimates based on taxonomic collections, reconciling an order of magnitude discrepancy between extrapolations of global diversity based on ecological samples of tropical communities with those based on sampling regional faunas.     

Host specificity of Lepidoptera in tropical and temperate forests

For numerous taxa, species richness is much higher in tropical than in temperate zone habitats. A major challenge in community ecology and evolutionary biogeography is to reveal the mechanisms underlying these differences. For herbivorous insects, one such mechanism leading to an increased number of species in a given locale could be increased ecological specialization, resulting in a greater proportion of insect species occupying narrow niches within a community. We tested this hypothesis by comparing host specialization in larval Lepidoptera (moths and butterflies) at eight different New World forest sites ranging in latitude from 15 degrees S to 55 degrees N. Here we show that larval diets of tropical Lepidoptera are more specialized than those of their temperate forest counterparts: tropical species on average feed on fewer plant species, genera and families than do temperate caterpillars. This result holds true whether calculated per lepidopteran family or for a caterpillar assemblage as a whole. As a result, there is greater turnover in caterpillar species composition (greater beta diversity) between tree species in tropical faunas than in temperate faunas. We suggest that greater specialization in tropical faunas is the result of differences in trophic interactions; for example, there are more distinct plant secondary chemical profiles from one tree species to the next in tropical forests than in temperate forests as well as more diverse and chronic pressures from natural enemy communities.     

Caterpillar-induced nocturnal plant volatiles repel conspecific females

Plants respond to insect herbivory by synthesizing and releasing complex blends of volatile compounds, which provide important host-location cues for insects that are natural enemies of herbivores. The effects of these volatile blends on herbivore behaviour have been investigated to only a limited extent, in part because of the assumption that herbivore-induced volatile emissions occur mainly during the light phase of the photoperiod. Because many moths-whose larvae are some of the most important insect herbivores-are nocturnal, herbivore-induced plant volatiles have not hitherto been considered to be temporally available as host-location cues for ovipositing females. Here we present chemical and behavioural assays showing that tobacco plants (Nicotiana tabacum) release herbivore-induced volatiles during both night and day. Moreover, several volatile compounds are released exclusively at night and are highly repellent to female moths (Heliothis virescens). The demonstration that tobacco plants release temporally different volatile blends and that lepidopteran herbivores use induced plant signals released during the dark phase to choose sites for oviposition adds a new dimension to our understanding of the role of chemical cues in mediating tritrophic interactions.     

Symbiotic fungal endophytes control insect host-parasite interaction webs

Symbiotic microorganisms that live intimately associated with terrestrial plants affect both the quantity and quality of resources, and thus the energy supply to consumer populations at higher levels in the food chain. Empirical evidence on resource limitation of food webs points to primary productivity as a major determinant of consumer abundance and trophic structure. Prey quality plays a critical role in community regulation. Plants infected by endophytic fungi are known to be chemically protected against herbivore consumption. However, the influence of this microbe-plant association on multi-trophic interactions remains largely unexplored. Here we present the effects of fungal endophytes on insect food webs that reflect limited energy transfer to consumers as a result of low plant quality, rather than low productivity. Herbivore-parasite webs on endophyte-free grasses show enhanced insect abundance at alternate trophic levels, higher rates of parasitism, and increased dominance by a few trophic links. These results mirror predicted effects of increased productivity on food-web dynamics. Thus 'hidden' microbial symbionts can have community-wide impacts on the pattern and strength of resource-consumer interactions.     

Global environmental controls of diversity in large herbivores

Large mammalian herbivores occupy half of the earth's land surface and are important both ecologically and economically, but their diversity is threatened by human activities. We investigated how the diversity of large herbivores changes across gradients of global precipitation and soil fertility. Here we show that more plant-available moisture reduces the nutrient content of plants but increases productivity, whereas more plant-available nutrients increase both of these factors. Because larger herbivore species tolerate lower plant nutrient content but require greater plant abundance, the highest potential herbivore diversity should occur in locations with intermediate moisture and high nutrients. These areas are dry enough to yield high quality plants and support smaller herbivores, but productive enough to support larger herbivores. These predictions fit with observed patterns of body size and diversity for large mammalian herbivores in North America, Africa and Australia, and yield a global map of regions with potentially high herbivore diversity. Thus, gradients of precipitation, temperature and soil fertility might explain the global distribution of large herbivore diversity and help to identify crucial areas for conservation and restoration.     

Jasmonate and salicylate induce expression of herbivore cytochrome P450 genes

Jasmonate and salicylate are plant-produced signals that activate plant defence genes after herbivory or pathogen attack. Amplification of these signals, evoked by either enemy attack or experimental manipulation, leads to an increase in the synthesis of toxic compounds (allelochemicals) and defence proteins in the plants. Although the jasmonate and salicylate signal cascades activate different sets of plant defence genes, or even act antagonistically, there is substantial communication between the pathways. Jasmonate and salicylate also contribute to protecting plants against herbivores by causing plants that experience insect damage to increase their production of volatile molecules that attract natural enemies of herbivorous insects. In response to plant defences, herbivores increase their production of enzymes that detoxify allelochemicals, including cytochrome P450s (refs 15, 16). But herbivores are potentially vulnerable to toxic allelochemicals in the duration between ingesting toxins and induction of detoxification systems. Here we show that the corn earworm Helicoverpa zea uses jasmonate and salicylate to activate four of its cytochrome P450 genes that are associated with detoxification either before or concomitantly with the biosynthesis of allelochemicals. This ability to 'eavesdrop' on plant defence signals protects H. zea against toxins produced by host plants.     

Tree use by koalas in a chemically complex landscape

Although defence against herbivores is often argued to be the main action of plant secondary metabolites (PSMs), very few examples have demonstrated that intraspecific variation in PSM concentrations influences foraging by wild vertebrate herbivores. Experiments with captive animals often indicate that PSM concentrations influence how much herbivores eat from individual plants, but these experiments do not replicate the subtle trade-offs in diet selection faced by wild animals, which must avoid predators and extremes of weather, interact with conspecifics, and achieve a balanced, nutritious diet, while avoiding intoxication by PSMs. We characterized the foliar chemistry of every tree from two Eucalyptus species available to a population of koalas (Phascolarctos cinereus) and considered rates of tree visitation over a ten-year period. We show that visitation rate was most strongly influenced by tree size, but that koalas also visited trees less frequently if the foliage contained either high concentrations of deterrent PSMs known as formylated phloroglucinol compounds, or low concentrations of nitrogen. Consequently, plant chemistry restricts the use of trees by this herbivore, and thus limits the food available to koalas and potentially influences koala populations.     

Evolutionary change from induced to constitutive expression of an indirect plant resistance

Induced plant resistance traits are expressed in response to attack and occur throughout the plant kingdom. Despite their general occurrence, the evolution of such resistances has rarely been investigated. Here we report that extrafloral nectar, a usually inducible trait, is constitutively secreted by Central American Acacia species that are obligately inhabited by ants. Extrafloral nectar is secreted as an indirect resistance, attracting ants that defend plants against herbivores. Leaf damage induces extrafloral nectar secretion in several plant species; among these are various Acacia species and other Fabaceae investigated here. In contrast, Acacia species obligately inhabited by symbiotic ants nourish these ants by secreting extrafloral nectar constitutively at high rates that are not affected by leaf damage. The phylogeny of the genus Acacia and closely related genera indicate that the inducibility of extrafloral nectar is the plesiomorphic or 'original' state, whereas the constitutive extrafloral nectar flow is derived within Acacia. A constitutive resistance trait has evolved from an inducible one, obviously in response to particular functional demands.     

Parasitic plants indirectly regulate below-ground properties in grassland ecosystems

Parasitic plants are one of the most ubiquitous groups of generalist parasites in both natural and managed ecosystems, with over 3,000 known species worldwide. Although much is known about how parasitic plants influence host performance, their role as drivers of community- and ecosystem-level properties remains largely unexplored. Parasitic plants have the potential to influence directly the productivity and structure of plant communities because they cause harm to particular host plants, indirectly increasing the competitive status of non-host species. Such parasite-driven above-ground effects might also have important indirect consequences through altering the quantity and quality of resources that enter soil, thereby affecting the activity of decomposer organisms. Here we show in model grassland communities that the parasitic plant Rhinanthus minor, which occurs widely throughout Europe and North America, has strong direct effects on above-ground community properties, increasing plant diversity and reducing productivity. We also show that these direct effects of R. minor on the plant community have marked indirect effects on below-ground properties, ultimately increasing rates of nitrogen cycling. Our study provides evidence that parasitic plants act as a major driver of both above-ground and below-ground properties of grassland ecosystems.     

Coevolution in multidimensional trait space favours escape from parasites and pathogens

Almost all species are subject to continuous attack by parasites and pathogens. Because parasites and pathogens tend to have shorter generation times and often experience stronger selection due to interaction than their victims do, it is frequently argued that they should evolve more rapidly and thus maintain an advantage in the evolutionary race between defence and counter-defence. This prediction generates an apparent paradox: how do victim species survive and even thrive in the face of a continuous onslaught of more rapidly evolving enemies? One potential explanation is that defence is physiologically, mechanically or behaviourally easier than attack, so that evolution is less constrained for victims than for parasites or pathogens. Another possible explanation is that parasites and pathogens have enemies themselves and that victim species persist because parasites and pathogens are regulated from the top down and thus generally have only modest demographic impacts on victim populations. Here we explore a third possibility: that victim species are not as evolutionarily impotent as conventional wisdom holds, but instead have unique evolutionary advantages that help to level the playing field. We use quantitative genetic analysis and individual-based simulations to show that victims can achieve such an advantage when coevolution involves multiple traits in both the host and the parasite.     

膝芒寄蝇族昆虫(双翅目:寄蝇科)研究进展

膝芒寄蝇族昆虫属微卵型,其幼虫主要寄生在鳞翅目、鞘翅目、膜翅目和双翅目的幼虫体内;通过触觉和嗅觉寻找寄主;寄生成功率较低,因此产卵量高,导致本族属阶元多,但种类少;寄蝇多样性分布与雌蝇产卵战略相关。世界已知本族88属,中国记录有49属139种。综述了膝芒寄蝇族的研究史和现状,介绍了其系统地位、形态分类、动物区系与分布、系统发育与演化的单系性、寄生生物学与害虫控制等方面的研究进展。分析了目前该类群存在的问题并展望了今后研究工作内容和方向。     

Low beta diversity of herbivorous insects in tropical forests

Recent advances in understanding insect communities in tropical forests have contributed little to our knowledge of large-scale patterns of insect diversity, because incomplete taxonomic knowledge of many tropical species hinders the mapping of their distribution records. This impedes an understanding of global biodiversity patterns and explains why tropical insects are under-represented in conservation biology. Our study of approximately 500 species from three herbivorous guilds feeding on foliage (caterpillars, Lepidoptera), wood (ambrosia beetles, Coleoptera) and fruit (fruitflies, Diptera) found a low rate of change in species composition (beta diversity) across 75,000 square kilometres of contiguous lowland rainforest in Papua New Guinea, as most species were widely distributed. For caterpillars feeding on large plant genera, most species fed on multiple host species, so that even locally restricted plant species did not support endemic herbivores. Large plant genera represented a continuously distributed resource easily colonized by moths and butterflies over hundreds of kilometres. Low beta diversity was also documented in groups with differing host specificity (fruitflies and ambrosia beetles), suggesting that dispersal limitation does not have a substantial role in shaping the distribution of insect species in New Guinea lowland rainforests. Similar patterns of low beta diversity can be expected in other tropical lowland rainforests, as they are typically situated in the extensive low basins of major tropical rivers similar to the Sepik-Ramu region of New Guinea studied here.     

Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment

Biodiversity is rapidly declining, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above- and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades.     

Role of ethylene signaling in the production of rice volatiles induced by the rice brown planthopper Nilaparvata lugens

In response to herbivory, plants release volatiles to attract the natural enemies of herbivores[1,2]. This phe- nomenon has been reported in more than 23 plant spe- cies[3―6], and several field studies have shown that these herbivore-induced volatiles en…     

高温胁迫对植食性昆虫影响研究进展

昆虫作为变温动物类群,极易遭受高温胁迫的影响。研究高温胁迫对植食性昆虫的影响,可为气候变暖背景下农林害虫的种群动态监测及地理适生区预测提供重要理论依据。笔者从植食性昆虫个体、种间互作关系、种群及群落3个层面,综述了高温胁迫对植食性昆虫影响的国内外进展。研究发现,高温胁迫会对植食性昆虫个体的生长发育、繁殖及生理生化带来负面效应,但昆虫自身也会进化出一系列基于形态、行为及基因表达的热胁迫响应机制。高温胁迫对寄主植物、天敌、共生微生物的影响也能够通过种间互作传递给植食性昆虫,并导致寄主植物-植食性昆虫-天敌3级营养关系发生变化。同时,高温胁迫还能通过影响昆虫个体及种间关系,间接影响昆虫的种群动态、种群多样性及其生态功能,并可能导致植食性昆虫的种群大爆发或衰退。最后,笔者认为此领域未来研究方向为:在个体层面上,应优化高温胁迫处理方式并综合考虑干旱和降水等环境因子,开展农林害虫种群的长期野外监测,应注重高温胁迫对昆虫参与生长发育繁殖功能基因的影响;在种间关系层面上,应关注高温胁迫对昆虫关联的复杂多样的食物网及互作体系的影响;此外,应结合高温胁迫对不同昆虫种类自身生理特性和行为模式的影响,阐明温度升高对昆虫群落的影响。     

Positive interactions among alpine plants increase with stress

Plants can have positive effects on each other. For example, the accumulation of nutrients, provision of shade, amelioration of disturbance, or protection from herbivores by some species can enhance the performance of neighbouring species. Thus the notion that the distributions and abundances of plant species are independent of other species may be inadequate as a theoretical underpinning for understanding species coexistence and diversity. But there have been no large-scale experiments designed to examine the generality of positive interactions in plant communities and their importance relative to competition. Here we show that the biomass, growth and reproduction of alpine plant species are higher when other plants are nearby. In an experiment conducted in subalpine and alpine plant communities with 115 species in 11 different mountain ranges, we find that competition generally, but not exclusively, dominates interactions at lower elevations where conditions are less physically stressful. In contrast, at high elevations where abiotic stress is high the interactions among plants are predominantly positive. Furthermore, across all high and low sites positive interactions are more important at sites with low temperatures in the early summer, but competition prevails at warmer sites.