Biodiversity of galling insects: historical, community and habitat effects in four neotropical savannas

Five hypotheses were tested to explain the pattern of galling insect species richness in four neotropical savanna physiognomies, 'canga ', 'campo sujo', 'cerrado' s. st., and 'cerradão', that occur in Minas Gerais, southeastern Brazil. We found 125 species of galling insects on 80 host plant species. The increase of plant species richness explained 35% of the variation in galling insect richness, corroborating the plant species richness hypothesis. Most of the galling species occurred on trees, followed by shrubs, and herbs. However, the difference in mean number of galls was only statistically significant between herbs and trees, corroborating partially the plant structural complexity hypothesis. A significant relationship was observed between galling species richness and density of herbs, and shrubs, corroborating partially the resource concentration hypothesis. Galling insect richness showed a negative correlation with magnesium, potassium, and zinc on soil, corroborating the soil fertility hypothesis. The content of magnesium, potassium, iron and CTC (T) explained 72% of the variation in galling insect richness. Plant family size positively influenced galling insect richness, corroborating the plant family size hypothesis. Overall, the results corroborate the hypothesis that predicts that habitat stress is the main factor generating the patterns of galling insect richness in Brazilian savannas.     

A meta-analysis of the effects of galling insects on host plant secondary metabolites

The idea that galling insects actively manipulate host plant chemistry has been previously documented but has not been quantified across a range of galler and host plant taxa. We present the first quantitative review of the relationship between insect galling and levels of secondary metabolites in host plants. Using meta-analytic techniques, we examined this relationship across 40 galler and host plant species combinations. We found that galling insects are associated with significantly higher levels of tannins and phenolics; however, no difference was found for volatiles. Hymenoptera, Diptera and Hemiptera were associated with higher levels of secondary metabolites; however, only Hymenoptera was significant. The climatic zone of the study area did not explain significant differences in gall-induced secondary metabolites. Overall the results show that the ability of galling insects to manipulate host plant secondary chemistry is widespread across insect and plant taxa. The evolutionary success of galling insects may be in part due to this unique ability.     

Global patterns in local number of insect galling species

Abstract. We evaluate a three-part hypothesis explaining why gall-inducing insect species richness is so high in scleromorphic vegetation: (1) persistence of low nutrient status scleromorphic leaves facilitates the galling habit in warm temperate latitudes; (2) favourable colonization sites for gallers result from reduced hygrothermal stress, high phenolics in the outer cortex of the gall, and reduced carnivore and fungal attack in the gall; and (3) in more mesic sites, mortality is high due to carnivore attack and invasion of galls by fungi. Over 280 samples of local species of galling herbivorous insects from fourteen countries on all continents except Antarctica revealed a strong pattern of highest richness in warm temperate latitudes, or their altitudinal equivalents. The peak of galling species richness on the latitudinal gradient from the equator into the Arctic was between 25 to 38° N or S. Galling species were particularly diverse in sclerophyllous vegetation, which commonly had greater than twelve species per local sample. In mesic, non-sclerophyllous vegetation types the number of galling species was lower with twelve or fewer species present. Many sites in sclerophyllous vegetation supported between thirteen and forty-six galling species locally, including campina islands in Amazonia, cerrado savanna in central Brazil, the Sonoran Desert in Arizona and Mexico, shrubland in Israel, fynbos in South Africa and coastal scleromorphic vegetation in Australia. At the same latitude, or its elevational equivalent, galling species richness was significantly higher in relatively xeric sites when compared to riparian or otherwise mesic habitats, even when scleromorphic vegetation dominated the mesic sites. The results were consistent with the hypothesis and extend to a more general level the patterns and predictions on the biogeography of gall-inducing insects.     

Vigour of a dioecious shrub and attack by a galling herbivore

1. The pattern of attack by the leaf‐galling insect Neopelma baccharidis (Homoptera: Psyllidae) was studied in three populations of the dioecious shrub Baccharis dracunculifolia (Asteraceae) in south‐eastern Brazil. The plant vigour hypothesis, which predicts higher rates of attack and increased herbivore performance on the longest plant shoots, was tested. This work also provides further information for the study of differential herbivory in dioecious plants. 2. In total, 9200 shoots were collected randomly from 46 male and 47 female plants belonging to the three populations. Shoot length, number of leaves per shoot, rate of galling, and survival of psyllids did not differ between male and female plants. Another population on the Campus of the Federal University of Minas Gerais was used only to determine the pattern of shoot growth. 3. The hypothesis of sex‐mediated herbivory was not corroborated in this study. 4. The frequency of galling increased with increasing shoot length, as predicted by the plant vigour hypothesis. Nevertheless, the number of oviposition sites (leaf buds) increased with shoot length. 5. The performance of the galling herbivore was not related to shoot length in the plant populations studied. 6. In conclusion, Neopelma baccharidis did not select shoots based on length only.     

Effects of plant architecture and water velocity on sediment retention by submerged macrophytes

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Host plant effects on the development and survivorship of the galling insect Neopelma baccharidis (Homoptera: Psyllidae)

In this study, the mortality factors acting upon the galling psyllid Neopelma baccharidis Burckhardt (Homoptera) caused by its host plant, Baccharis dracunculifolia De Candole (Asteraceae) were analysed. In March 1999, 982 galls of the same cohort were randomly marked on 109 individuals of B. dracunculifolia in the field. Galls were censused each month during their development, from April to August, and dead galls were collected and analysed for mortality factors. Gall dehiscence rates were calculated for each month. The major mortality source of N. baccharidis was gall dropping (13.2% of the original cohort), which is probably a normal outcome of previous mortality caused by the other factors observed in this study. Unknown factors killed 11.7% of this gall population and were ascribed to plant resistance during gall development. Empty galls represented 7.7% of the observed mortality and may be a consequence of egg retention or egg mortality/abortion related to variations in plant quality. Shoot mortality was high during the dry season and killed 7.5% of the galls, but this impact was minimized after the third month from gall formation due to the ability of nymphs to accelerate development and emerge from galls on dying shoots. However, the size of dehisced galls on dead shoots tended to be smaller, possibly affecting adult performance. Mortality of N. baccharidis attributed to B. dracunculifolia strongly controlled the galling insect population, killing 40.7% of the original cohort of galls. Plant‐mediated mortality was caused by often neglected factors acting predominantly during the first 3 months of development, which are critical to gall survivorship. These results reinforce the importance of bottom‐up forces in plant‐insect systems.     

虫瘿—昆虫与植物互作的奇特产物

虫瘿是昆虫刺激产生的植物不正常组织。造瘿昆虫几乎能在高等植物的所有类群中产生形态各异的虫瘿,甚至能在同一种植物上形成多种形态不一的虫瘿。虫瘿形成的机制仍不清楚,普遍认为是昆虫控制了虫瘿的形成,但植物细胞也参与了虫瘿形成的调控。为了探索这一神秘的领域,科学家们已经对昆虫的刺激物以及植物的反应作了大量的研究,未来工作的难点将是如何把二者联系起来。     

The effect of plant density on epiphytic macroinvertebrates associated with a submerged macrophyte,Lagarosiphon ilicifolius Obermeyer,in Lake Kariba,Zimbabwe

The effect of variations in the density of a submerged macrophyte, Lagarosiphon ilicifolius, on epiphytic macroinvertebrate community structure in the shallow waters of a sheltered bay of Lake Kariba were investigated. The body size class distributions of a mayfly, Cloeon (Ephemeroptera: Baetidae), and the damselfly family, Coenagrionidae, were also assessed with respect to variation in vegetation density. Macroinvertebrates were sampled from low-, moderate- and high-density beds of L. ilicifolius. There were no significant differences in individual taxon and total macroinvertebrate abundances, macroinvertebrate richness and diversity with respect to vegetation density. In all three density categories the functional feeding group (FFG) composition was dominated by collector-gatherers and collector-filterers. The abundance of the two FFGs did not change significantly within each, as well as among, the three vegetation-density categories (ANOVA, p > 0.05). The largest size class of Cloeon occurred only in high-density beds, whereas the largest coenagrionid individuals were obtained from low- and moderate-density beds and were absent from high-density beds. The results suggest that variation in the density of Lagarosiphon does not affect epiphytic macroinvertebrate community structure, but does affect body-size distributions of macroinvertebrate taxa, probably by affecting predator-prey interactions.     

The effect of herbivore density on host plant mediated interactions between two insects

Recent work suggests that foliar- and root-feeding insects can interact in a novel plus-minus fashion. However, the effects of herbivores may be different at different densities. This paper describes two laboratory experiments investigating the effect of increasing insect density on plant performance and the host-plant mediated interaction between a leaf-mining fly and root-feeding chafer larvae. Above- and below-ground insect herbivory decreased plant performance. The effects of the root feeder were, for some parameters, different between the two densities of chafer larvae. Leaf mining significantly decreased the performance of the chafer larvae, while root herbivory was found to increase the pupal weight (related to fecundity) of the leaf miner. The effects of root herbivory suggest that the relationships between herbivore density and plant performance may be curved as the greater chafer density had no significant increased effect on the plants or foliar-feeding insects.     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

Impact in quarantine of the galling weevil Lepidapion argentatum on shoot growth of French broom (Genista monspessulana), an invasive weed in the western U.S.

In weed biocontrol, there is a need for pre-release efficacy assessments for potential agents. Genista monspessulana ((L.) L. A. S. Johnson (Fabaceae), French broom) is an invasive perennial shrub in the western U.S. The galling weevil Lepidapion argentatum Gerstaecker is a potential biocontrol agent. The impact of increasing weevil density on galling damage, plant height, width, leaf damage, and relative growth rate (RGR) was assessed in greenhouse experiments on two to three-month-old seedlings infested with either one or three weevils. Infestation by three female weevils caused 48% more galls producing 27% more larvae than did infestation with one female while causing only 1% leaf damage and no difference in total leaf area. Infestation with multiple weevils caused a 55% and 29% decrease in plant height and canopy width respectively, while single-weevil infestation decreased height by 32% and width to the same degree as for multiple weevils. The RGR of seedlings infested with three weevils was three times slower than the controls, while growth was reduced 2-fold by single-weevil infestation. Reductions in plant size and growth rate induced by weevil galling could reduce plant competitive survival to reproduction and also plant population dispersal as seedlings. Our results suggest that L. argentatum has the potential to cause impact to French broom seedlings if released in the invasive range.     

The importance of plant relatedness for host utilization among phytophagous insects

Current methods for measuring similarity among phytophagous insect communities fail to consider the phylogenetic relationship between host plants. We analysed this relation based on 3580 host observations of 1174 beetle species associated with 100 species of angiosperms in two different forest types in Panama. We quantified the significance of genetic distance as well as taxonomic rank among angiosperms in relation to species overlap in beetle assemblages. A logarithmic model describing the decrease in beetle species similarity between host-plant species of increasing phylogenetic distance explains 35% of the variation. Applied to taxonomic rank categories the results imply that except for the ancient branching of monocots from dicots, only adaptive radiations of plants on the family and genus level are important for host utilization among phytophagous beetles. These findings enable improvements in estimating host specificity and species richness through correction for phylogenetic relatedness between hosts and consideration of the host-specific fauna associated with monocots.     

Effects of altered precipitation on insect community composition and structure in a meadow steppe

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Effects of plant diversity and structural complexity on parasitoid behaviour in a field experiment

1. In natural ecosystems, plants containing hosts for parasitoids are often embedded within heterogeneous plant communities. These plant communities surrounding host‐infested plants may influence the host‐finding ability of parasitoids. 2. A release‐recapture‐approach was used to examine whether the diversity and structural complexity of the community surrounding a host‐infested plant influences the aggregation behaviour of the leaf‐miner parasitoid Dacnusa sibirica Telenga and naturally occurring local leaf‐miner parasitoids. Released and locally present parasitoids were collected on potted Jacobaea vulgaris Gaertn.plants infested with the generalist leaf‐miner Chromatomyia syngenesiae Hardy. The plants were placed in experimentally established plant communities differing in plant diversity (1–9 species) and habitat complexity (bare ground, mown vegetation, and tall vegetation). Additionally, parasitoids were reared out from host mines on the trap plants. 3. Plant diversity did not influence the mean number of recaptured D. sibirica or captures of other locally present parasitoids but the number of recaptured parasitoids was influenced by habitat complexity. No D. sibirica parasitoids were recaptured in the bare ground plots or plots with mown vegetation. The mean number of recaptured D. sibirica generally increased with increasing complexity of the plant community, whereas locally present parasitoids were captured more frequently in communities with more bare ground. There was a unimodal relationship between the number of reared out parasitoids and diversity of the surrounding vegetation with the highest density of emerged parasitoids at intermediate diversity levels. 4. The present study adds to the thus far limited body of literature examining the aggregation behaviour of parasitoids in the field and suggests that the preference of parasitoids to aggregate in complex versus simple vegetation is association specific and thus depends on the parasitoid species as well as the identity of the plant community.     

Effect of physiological status on olfactory and visual responses of female Hypothenemus hampei during host plant colonization

The response of Hypothenemus hampei (Ferr.) (Coleoptera, Scolytidae) (coffee berry borer, CBB) females to olfactory stimuli was demonstrated to be related with the physiological status; especially with age, whether they were mated or not, and whether they have already oviposited. A special behaviour, the abandonment of the native berry, was also found to play a major role. In contrast, response to visual stimuli (false coffee berries) was found to be independent of the physiological status of CBB females. Variations in behavioural responses to visual and olfactory stimuli are discussed in relation to colonization strategy.     

Determinants of growth rate on chemically heterogeneous host plants by specialist insects

Nine Passiflora species growing sympatrically in a Costa Rican rain forest were screened for the presence of alkaloids, tannins, cyanogenic compounds, saponins and cardenolides. The first three chemical classes were found to be present and the Passiflora species could be classified into five groups according to the presence/absence of each chemical class. In order to assess the effects of this chemical variability on specialist herbivores, larvae of Heliconius ismenius and H. melpomane were experimentally introduced to seven species of Passiflora host plants growing naturally in the rain forest. Survival and growth rates were subsequently monitored at 2 day intervals and were found to be independent of each other. Larval growth rate and larval survivorship were not significantly affected by cyanogenesis in the host plants, nor by the presence of alkaloids, tannins and non-tanning phenolic compounds. Larval growth rate was found to be correlated to percentage nitrogen in the host plants, but was independent of host water content and non-structural carbohydrate content. Although these insects would be classified as host-specific by most authors, the results indicate the presence of a sufficiently general detoxication apparatus to consume many different classes of potential toxins with no apparent ill effects. We conclude that Passiflora natural product diversity does not cause major deleterious effects in Heliconius growth characteristics and that these effects are not the major component maintaining the current pattern of larval feeding specialization in this plant-herbivore system.     

Aquatic macrophytes, macroinvertebrate associations and water levels in a lowland Tasmanian river

Aquatic macrophytes are a common habitat for macroinvertebrates and may occupy depth zones in the littoral region of lowland rivers. Studies have indicated that different species of macrophyte typically support different assemblages, abundances and numbers of species of macroinvertebrates. This has often been attributed to differences in the dissectedness of stems and leaves of the macrophytes, resulting in differences in the surface area and/or the number of microhabitats available to invertebrates. I set out to measure the abundance and taxonomic richness and to describe the macroinvertebrate assemblages associated with three species of aquatic macrophyte in a pool in the Macquarie River, Tasmania and to examine responses of these variables to changes in water levels over summer. The macrophyte species sampled wereMyriophyllum simulans/variifolium, Triglochin procera} and Eleocharis sphacelata, each one differing in the dissectedness of its stems and leaves and its location in the littoral zone. Whereas the greatest abundance of macroinvertebrates was found associated in all months (i.e. at all water levels) with the structurally complex and shallowest macrophyte species, Myriophyllum, the number of taxa associated with this species was in several cases lower than for the structurally simpler and deeper water Triglochin and Eleocharis. While water depth and total plant biomass of samples were often correlated with invertebrate abundance and richness, these relationships were different for each macrophyte species. Of the nine most common invertebrate taxa collected from all samples, the abundances of more than half showed consistent differences among macrophyte species across months, two showed differences among macrophytes, but with an interaction with month and two showed no differences among macrophytes. There were major differences in the invertebrate assemblages associated with each macrophyte species in any one month, however, there was also a large turnover of taxa associated with the species of macrophytes from one month to the next. Changes in water level and concomitant changes in environmental variables are suggested as factors influencing the invertebrate fauna in the littoral zone of the pool of the Macquarie River. It is thus important for river managers to be aware that species of macroinvertebrates are not evenly distributed across species of macrophyte and that water levels and their influence on macrophytes as invertebrate habitat may play an integral part in determining the abundance, richness and assemblage of invertebrates in rivers.     

Epidemiological prediction of the distribution of insects of medical significance: comparative distributions of fleas and sucking lice on the rat host Rattus norvegicus in Yunnan Province, China

Determining the distribution patterns of ectoparasites is important for predicting the spread of vector-borne diseases. A simple epidemiological model was used to compare the distributions of two different taxa of ectoparasitic insects, sucking lice (Insecta: Siphonaptera) and fleas (Insecta: Anoplura), on the same rodent host, Rattus norvegicus Berkenhout (Rodentia: Muridae), in Yunnan Province, China. Correlations between mean abundance and prevalence were determined. Both fleas and sucking lice were aggregated on their hosts, and sucking lice showed a higher degree of aggregation than fleas. The prevalence of both fleas and sucking lice increased with log-transformed mean abundance and a highly linear correlation and modelling efficiency of predicted prevalence against observed prevalence were obtained. The results demonstrate that prevalence can be explained simply by mean abundance.