Differences in meristems between monocots and dicots and susceptibility to attack by gall-inducing insects

We used comparative methods that account for the phylogenetic correlations among species to test hypotheses about the community of gall-inducing insects on dicotyledonous and monocotyledonous plants and woody and herbaceous angiosperms in the UK. We found that the species richness of gall-inducing insects on dicots was greater than on monocots and that the odds of a dicot having an associated gall-inducing insect is 42% higher than for a monocot. Woody angiosperms have higher species richness of associated gall-inducing insects than do herbaceous angiosperms. Furthermore, using a Monte Carlo analysis we found that attacks by gall-inducing insects on monocot families were phylogenetically clustered in the order Poales, particularly within the grass family Poaceae. We suggest that the higher risk of attack on dicots and higher species richness of gall-inducing insects on woody angiosperms, which are exclusively dicots, arises because of differences in the abundance or susceptibility of dicot meristems to attack by gall-inducing insects. Architectural and anatomical differences between monocots and dicots that give rise to differences in meristem abundance and anatomy appear to play an important role in determining the occurrence and richness of associated gall-inducing insects on host plants.     

Effects of non-native plants on the native insect community of Delaware

Due to the lack of a co-evolutionary history, the novel defenses presented by introduced plants may be insurmountable to many native insects. Accordingly, non-native plants are expected to support less insect biomass than native plants. Further, native insect specialists may be more affected by introduced plants than native generalist herbivores, resulting in decreased insect diversity on non-native plants due to the loss of specialists. To test these hypotheses, we used a common garden experiment to compare native insect biomass, species richness, and the proportion of native specialist to native generalist insects supported by 45 species of woody plants. Plants were classified into three groupings, with 10 replicates of each species: 15 species native to Delaware (Natives), 15 non-native species that were congeneric with a member of the Native group (Non-native Congeners), and 15 non-native species that did not have a congener present in the United States (Aliens). Native herbivorous insects were sampled in May, June, and July of 2004 and 2005. Overall, insect biomass was greater on Natives than Non-native Congeners and Aliens, but insect biomass varied unpredictably between congeneric pair members. Counter to expectations, Aliens held more insect biomass than did Non-native Congeners. There was no difference in species richness or the number of specialist and generalist species collected among the three plant groupings in either year, although our protocol was biased against sampling specialists. If these results generalize to other studies, loss of native insect biomass due to introduced plants may negatively affect higher trophic levels of the ecosystem.     

Comparing galling insect richness among Neotropical savannas: effects of plant richness, vegetation structure and super-host presence

Plant species diversity maintains the stability of ecosystems and the diversity of consumer species such as insect herbivores. Considering that gall-inducing insects are highly specialized on their host plants and dependent on the occurrence, abundance and distribution of plants, we evaluated the diversity patterns of gall-inducing insect along Brazilian Neotropical savannas and the potential role of plant species richness, vegetation structure and super-host presence on determining these patterns. We found 1,882 individual plants that belonged to 64 different host plant species grouped in 31 families, associated to 112 galling insect species. The galling richness was positively influenced by plant species richness and the presence of the super-host genus Qualea (Vochysiaceae). Plant species richness explained 48 % of the galling richness and areas with presence of super-hosts had more than twice of galling species than areas where they were absent. On the other hand, we found no evidence that larger plants hosted more species of galling insects. We observed that for the diversity of galling insects in the Brazilian Cerrado, vegetation structure explained almost the same portion as plant richness, because structural variables did not have an effect on residuals of galling richness and plant richness regression. Our findings suggests that plant richness has a more important role on the mitigation of natural enemies and adaptive radiation of galling species, while structural aspects of the vegetation does not seem to have that effect. Furthermore, we show that the super-host taxa provide an increment in local galling richness because they present a great diversity of local number of gall morphospecies (i.e. alpha diversity) and the high turnover of morphospecies among different localities (i.e. beta diversity). Therefore we argue that the quality of resources (richness and super host presence) appears to be a most important factor for the diversity of galling insects in Neotropical systems, than the amount of resources.     

Native plant diversity increases herbivory to non-natives

There is often an inverse relationship between the diversity of a plant community and the invasibility of that community by non-native plants. Native herbivores that colonize novel plants may contribute to diversity–invasibility relationships by limiting the relative success of non-native plants. Here, we show that, in large collections of non-native oak trees at sites across the USA, non-native oaks introduced to regions with greater oak species richness accumulated greater leaf damage than in regions with low oak richness. Underlying this trend was the ability of herbivores to exploit non-native plants that were close relatives to their native host. In diverse oak communities, non-native trees were on average more closely related to native trees and received greater leaf damage than those in depauperate oak communities. Because insect herbivores colonize non-native plants that are similar to their native hosts, in communities with greater native plant diversity, non-natives experience greater herbivory.     

Species richness of herbivores on exotic host plants increases with time since introduction of the host

Aim Species richness of insect herbivores feeding on exotic plants increases with abundance as well as range size of the host in the area of introduction. The formation of these herbivore assemblages requires a certain amount of time, and the richness of insect faunas should also increase with the length of time an exotic plant has been present in the introduced range. Location Central Europe. Methods We analysed the variation in species richness of leaf‐chewing Lepidoptera larvae and sap‐sucking Auchenorrhyncha (Hemiptera) associated with 103 exotic woody plant species in Germany in relation to time since introduction, range size, growth form (trees versus shrubs), biogeographical origin (distance from Central Europe) and taxonomic isolation of the host plant (presence or absence of a native congener in the introduced area). Results Using simple correlation analyses we found for Lepidoptera and Auchenorrhyncha that species richness increased with time since introduction of the host plant. For the Lepidoptera the increase of species richness with time since introduction remained significant even after removing the effects of all other independent variables. Main conclusions Our results provide some evidence that assemblages of insects on exotic plants do not reach saturation within a time scale of few hundred years. This contrasts with previous findings for crop plants.     

Host plant predictability and the feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores

Summary Host plant preferences for 34 insect herbivore species are reported. Most polyphagous herbivores feeding on annuals, herbaceous perennials, and woody perennials show distinct preferences for the least abundant plant species among their various host plants. In addition, some populations of widely distributed polyphagous species are much more specialized in their diet than host plant lists alone would suggest. The high level of polyphagy on annuals and herbaceous perennials is suggested to be strongly influenced by the unpredictability of the host plant that is, in turn, controlled by environmental variability. Oligophagous herbivores preferred the least abundant woody perennials on the study sites. Ten of the 22 monophagous herbivores preferred the rarest of all the plant species on the same sites.     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.     

Diversity and abundance patterns of phytophagous insect communities on alien and native host plants in the Brassicaceae

The herbivore load (abundance and species richness of herbivores) on alien plants is supposed to be one of the keys to understand the invasiveness of species. We investigate the phytophagous insect communities on cabbage plants (Brassicaceae) in Europe. We compare the communities of endophagous and ectophagous insects as well as of Coleoptera and Lepidoptera on native and alien cabbage plant species. Contrary to many other reports, we found no differences in the herbivore load between native and alien hosts. The majority of insect species attacked alien as well as native hosts. Across insect species, there was no difference in the patterns of host range on native and on alien hosts. Likewise the similarity of insect communities across pairs of host species was not different between natives and aliens. We conclude that the general similarity in the community patterns between native and alien cabbage plant species are due to the chemical characteristics of this plant family. All cabbage plants share glucosinolates. This may facilitate host switches from natives to aliens. Hence the presence of native congeners may influence invasiveness of alien plants.     

Gall‐forming insect species richness along a non‐scleromorphic vegetation rainfall gradient in South Africa: The importance of plant community composition

Abstract Currently there is no single accepted hypothesis to explain gall‐forming insect species richness at a particular locality. Hygrothermal stress, soil nutrient availability, plant species richness, plant structural complexity, plant family or genus size, and host plant geographical range size have all been implicated in the determination of gall‐forming insect species richness. Previous studies of such richness at xeric sites have included predominantly scleromorphic vegetation, usually on nutrient‐poor soils. This study is the first to investigate gall‐forming insect species richness of xeric, non‐scleromorphic vegetation. Two habitat types were sampled at each of five localities across a rainfall gradient in the savanna biome of South Africa. The habitat types differed with respect to plant species composition and topography. Gall‐forming insect species richness did not increase with increasing hygrothermal stress or decreasing soil fertility. Rather, gall‐forming insect species richness was largely dependent on the presence of Terminalia sericea as well as other members of the Combretaceae and Mimosaceae. Plots where all these taxa were present had the highest gall‐forming insect species richness, up to 15 species, whereas plots with none of these taxa had a maximum of four galling‐insect species. Despite herb, shrub and tree strata not differing in gall‐forming insect species richness, insect galls were more common on woody than non‐woody plants. Also, stem galls were more frequent than apical or leaf galls. An alternative hypothesis to explain local gall‐forming insect species richness is suggested: galling insects may preferentially select those plant species with characteristics such as chemical toxicity, mechanical strength, degree of lignification or longevity that can be manipulated to benefit the galler. Thus plant community composition should be considered when attempting to explain gall‐forming insect species richness patterns.     

Plant architecture and meristem dynamics as the mechanisms determining the diversity of gall-inducing insects

Plant architecture is considered to affect herbivory intensity, but it is one of the least studied factors in plant–insect interactions, especially for gall-inducing insects. This study aimed to investigate the influence of plant architecture on the speciose fauna of gall-inducing insects associated with 17 species of Baccharis. Five architectural variables were evaluated: plant height, number of fourth-level shoots, biomass, average level and number of ramifications. The number of galling species associated with each host plant species was also determined. To test the effects of plant architecture on gall richness at the individual level, we used another data set where the number of fourth-level shoots and gall richness were determined for B. concinna, B. dracunculifolia, and B. ramosissima every 3 weeks during 1 year. The average similarity between host species based on gall fauna was low (9%), but plants with the same architectural pattern tended to support similar gall communities. The most important architectural trait influencing gall richness at the species level was the number of fourth-level shoots, which is indicative of the availability of plant meristems, a fundamental tissue for gall induction and development. This variable also showed a positive correlation with gall richness at the individual level. We propose that variations in gall richness among host species are driven by interspecific differences in plant architecture via availability of young, undifferentiated tissue, which is genetically controlled by the strength of the apical dominance. Plant architecture should have evolutionary consequences for gall communities, promoting insect radiation among architecturally similar plants through host shift and sympatric speciation. We also discuss the role of plant architecture in the global biogeography of gall-inducing insects. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Contrasting patterns of herbivore and predator pressure on invasive and native plants

Invasive non-native plant species often harbor fewer herbivorous insects than related native plant species. However, little is known about how herbivorous insects on non-native plants are exposed to carnivorous insects, and even less is known on plants that have recently expanded their ranges within continents due to climate warming. In this study we examine the herbivore load (herbivore biomass per plant biomass), predator load (predator biomass per plant biomass) and predator pressure (predator biomass per herbivore biomass) on an inter-continental non-native and an intra-continental range-expanding plant species and two congeneric native species. All four plant species co-occur in riparian habitat in north-western Europe. Insects were collected in early, mid and late summer from three populations of all four species. Before counting and weighing the insects were classified to trophic guild as carnivores (predators), herbivores, and transients. Herbivores were further subdivided into leaf-miners, sap-feeders, chewers and gallers. Total herbivore loads were smaller on inter-continental non-native and intra-continental range-expanding plants than on the congeneric natives. However, the differences depended on time within growing season, as well as on the feeding guild of the herbivore. Although the predator load on non-native plants was not larger than on natives, both non-native plant species had greater predator pressure on the herbivores than the natives. We conclude that both these non-native plant species have better bottom-up as well as top-down control of herbivores, but that effects depend on time within growing season and (for the herbivore load) on herbivore feeding guild. Therefore, when evaluating insects on non-native plants, variation within season and differences among feeding guilds need to be taken into account.     

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and micro-habitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having a geographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system.     

Misleading herbivory in a tropical tree

The damage caused by herbivores can be confused with the drop of necrotic spots due to hypersensitive reactions. The negative impact of sessile herbivores on host plants is minimized by hypersensitive reactions that result in the death of attacked tissue. We reported a phenomenon that remarkably resembles herbivore damage, but is in fact a reaction to endophytic herbivores or pathogens. Although the damages on Tapirira guianensis leaves (Anacardiaceae) appear to be chew marks from herbivores, they are in fact dropped necrotic spots caused by gall-inducing insects. The phenomenon is widespread and challenges the view that gall-inducing insects inflict less damage on host plants, which was previously stated according to the rates of herbivory inflicted by free-feeding herbivores. The study highlights the need to reassess the past evaluations for some plant species, as they may have overestimated herbivory rates.     

Pattern and process in the evolution of insect-plant associations: Yponomeuta as an example

Phylogenetic studies are increasing our understanding of the evolution of associations between phytophagous insects and their host plants. Sequential evolution, i.e. the shift of insect herbivores onto pre-existing plant species, appears to be much more common than coevolution, where reciprocal selection between interacting insects and plants is thought to induce chemical diversification and resistance in plants and food specialization in insects.Extreme host specificity is common in phytophagous insects and future studies are likely to reveal even more specialization. Hypotheses that assume that food specialists have selective advantages over generalists do not seem to provide a general explanation for the ubiquity of specialist insect herbivores. Specialists are probably committed to remain so, because they have little evolutionary opportunity to reverse the process due to genetically determined constraints on the evolution of their physiology or nervous system. The same constraints might result in phylogenetic conservatism, i.e. the frequent association of related insect herbivores with related plants. Current phylogenetic evidence, however, indicates that there is no intrinsic direction to the evolution of specialization.Historical aspects of insect-host plant associations will be illustrated with the small ermine moth genus Yponomeuta. Small ermine moths show an ancestral host association with the family Celastraceae. The genus seems to be committed to specialization per se rather than to a particular group of plants. Whatever host shift they have made in their evolutionary past (onto Rosaceae, Crassulaceae, and Salicaceae), they remain monophagous. The oligophagous Y. padellus is the only exception. This species might comprise a mosaic of genetically divergent host-associated populations.     

Gall-induction in insects: evolutionary dead-end or speciation driver?

Background

The tree of life is significantly asymmetrical - a result of differential speciation and extinction - but general causes of such asymmetry are unclear. Differences in niche partitioning are thought to be one possible general explanation. Ecological specialization might lead to increases in diversification rate or, alternatively, specialization might limit the evolutionary potential of specialist lineages and increase their extinction risk. Here we compare the diversification rates of gall-inducing and non-galling insect lineages. Compared with other insect herbivores feeding on the same host plant, gall-inducing insects feed on plant tissue that is more nutritious and less defended, and they do so in a favorable microhabitat that may also provide some protection from natural enemies. We use sister-taxon comparisons to test whether gall-inducing lineages are more host-specific than non-galling lineages, and more or less diverse than non-gallers. We evaluate the significance of diversity bipartitions under Equal Rates Markov models, and use maximum likelihood model-fitting to test for shifts in diversification rates.

Results

We find that, although gall-inducing insect groups are more host-specific than their non-galling relatives, there is no general significant increase in diversification rate in gallers. However, gallers are found at both extremes - two gall-inducing lineages are exceptionally diverse (Euurina sawflies on Salicaceae and Apiomorpha scale insects on Eucalytpus), and one gall-inducing lineage is exceptionally species-poor (Maskellia armored scales on Eucalyptus).

Conclusions

The effect of ecological specialization on diversification rates is complex in the case of gall-inducing insects, but host range may be an important factor. When a gall-inducing lineage has a host range approximate to that of its non-galling sister, the gallers are more diverse. When the non-galler clade has a much wider host range than the galler, the non-galler is also much more diverse. There are also lineage-specific effects, with gallers on the same host group exhibiting very different diversities. No single general model explains the observed pattern.     

Allelopathic invasive tree (<Emphasis Type="Italic">Rhamnus cathartica</Emphasis>) alters native plant communities

Many plants release allelopathic chemicals that can inhibit germination, growth, and/or survival in neighboring plants. These impacts appear magnified with the invasion of some non-native plants which may produce allelochemicals against which native fauna have not co-evolved resistance. Our objective was to examine the potential allelopathic impact of an invasive non-native shrub/tree on multiple plant species using field observation and experimental allelopathy studies. We surveyed and collected an invasive, non-native tree/shrub (Rhamnus cathartica) at Tifft Nature Preserve (a 107-ha urban natural area near Lake Erie in Buffalo, NY). We also surveyed understory plant communities in the urban forest to examine correlations between R. cathartica abundance and local plant community abundance and richness. We then used experimental mesocosms to test if patterns observed in the field could be explained by adding increased dosages of R. cathartica to soils containing five plant species, including native and non-native woody and herbaceous species. In the highly invaded urban forest, we found that herbaceous cover, shrubs and woody seedlings negatively covaried with R. cathartica basal area and seedlings density. In the mesocosm experiments, R. cathartica resulted in significant decreases in plant community species richness, abundance, and shifted biomass allocation from roots. Our results provide evidence that R. cathartica is highly allelopathic in its invaded range, that R. cathartica roots have an allelopathic effect and that some plant species appear immune. We suggest that these effects may explain the plant’s ability to form dense monocultures and resist competitors, as well as shift community composition with species-specific impacts.     

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.     

Exotic herbivores and fire energy drive standing herbaceous biomass but do not alter compositional patterns in a semiarid savanna ecosystem

Questions

Fire regime alterations are pushing open ecosystems worldwide past tipping points where alternative steady states characterized by woody dominance prevail. This reduces the frequency and intensity of surface fires, further limiting their effectiveness for controlling cover of woody plants. In addition, grazing pressure (exotic or native grazers) can reinforce woody encroachment by potentially reducing fine-fuel loads. We investigated the effects of different fire energies on the herbaceous plant community, together with mammalian wildlife herbivory (exotic and native combined) exclusion, to inform best management practices.

Location

Texas semi-arid savanna, southern Great Plains, USA.

Methods

We conducted an experiment in which we manipulated fire intensity and herbivore access to herbaceous biomass in a split-plot design. We altered fire energy via fuel addition rather than applying fire under different environmental conditions to control for differences in standing biomass and composition attributable to differential plant physiological status and fire season.

Results

High-energy fire did not reduce herbaceous biomass or alter plant community composition, although it did increase among-plot variability in composition and forb biomass relative to low-energy fire and non-burned controls. Grazing pressure from native and non-native mammalian herbivores reduced above-ground herbaceous biomass regardless of fire treatments, but did not alter community composition.

Conclusions

Managers seeking to apply high-intensity prescribed fire to reduce woody encroachment will not negatively impact herbaceous plant productivity or alter community composition. However, they should be cognizant that repeated fires necessary for greatly reducing woody plants in heavily invaded areas might be difficult to accomplish due to fine-fuel reduction from wild herbivores. High fencing to restrict access by wildlife herbivores or culling might be necessary to build fuels sufficient to conduct high-intensity burns for woody-plant reduction.     

The Effect of Host-Plant Phylogenetic Isolation on Species Richness,Composition and Specialization of Insect Herbivores: A Comparison between Native and Exotic Hosts

Understanding the drivers of plant-insect interactions is still a key issue in terrestrial ecology. Here, we used 30 well-defined plant-herbivore assemblages to assess the effects of host plant phylogenetic isolation and origin (native vs. exotic) on the species richness, composition and specialization of the insect herbivore fauna on co-occurring plant species. We also tested for differences in such effects between assemblages composed exclusively of exophagous and endophagous herbivores. We found a consistent negative effect of the phylogenetic isolation of host plants on the richness, similarity and specialization of their insect herbivore faunas. Notably, except for Jaccard dissimilarity, the effect of phylogenetic isolation on the insect herbivore faunas did not vary between native and exotic plants. Our findings show that the phylogenetic isolation of host plants is a key factor that influences the richness, composition and specialization of their local herbivore faunas, regardless of the host plant origin.