Bracken, ants and extrafloral nectaries. III. How insect herbivores avoid ant predation

Abstract. 1. Ants (Myrmica spp. and Formica lemani) visiting the extrafloral nectaries of bracken, Pteridium aquilinum, imposed heavy mortality on caterpillars of a novel moth species experimentally introduced onto bracken fronds. However, the exclusion of ants from bracken fronds had no significant effect on adapted bracken-feeding herbivores in Britain.
2. The feeding stages of British bracken-feeding insects are immune to, or can largely avoid, predation by ants in a variety of ways. Gall formers and miners cannot be attacked by these ants. Two other species hide, one inside tied leaves, the other in a mass of 'spittle'. Another group of species jumps away from, or falls off the plant when touched by ants. Sawfly caterpillars in the genera Strongylogaster, Aneugmenus and Tenthredo have viscous, distasteful haemolymph that repels ants.
3. No species of bracken herbivore has an absolute temporal refuge from ants; during their development they all overlap with ants to some degree.
4. Ant predation appears to have played a significant role in determining the contemporary structure of British bracken-feeding insect communities. Distasteful haemolymph in sawfly caterpillars may have evolved in response to selection from ant predation. Other species may fortuitously possess characteristics, evolved in response to a variety of selective forces, that also reduce the impact of ants; without such characteristics, however, we postulate that they would be unable to live on this plant. An absence of external, foliage feeding Lepidoptera early in the spring, a high proportion of sawfly species, and a high proportion of gall-formers and miners may all be characteristics of the bracken herbivore community which have been influenced by ant predation.     

Herbivorous arthropods on bracken (Pteridium aquilinum (L.) Khun) in Australia compared with elsewhere

Abstract The herbivorous arthropod fauna of bracken fern (Pteridium aquilinum (L.) Khun), at a site near Sydney, Australia, is described and compared with previously reported bracken faunas in other geographic regions. Monthly sampling over 18 months found 17 species of herbivorous arthropods (15 insect and two mite species) from five orders. At the ordinal level, the mixture differed substantially from the bracken faunas of sites in Britain and Papua New Guinea. Notable was the presence of Thysanoptera and Acari, and the absence of Coleoptera and Hymenoptera. The mixtures of orders/families represented in the site bracken faunas in Britain, and less so in Australia, resembled those in the pool of herbivorous arthropods in those regions. Further, the mixture of orders on bracken was more similar to the mixture of orders on other ferns than to the mixture of orders among herbivorous insects on all plants; such similarity was not evident at family level. Compared with sites in other regions, the Sydney site had an abundance of pinna-sucking species and a dearth of mining species. Differences between regions in feeding niches most occupied tended to correspond with the differences in orders represented. Not all features of the fauna of bracken near Sydney reflected differences in the general herbivorous arthropod fauna of Australia compared with other regions, or differences between the herbivore faunas of ferns and seed plants. Its composition must be attributed in part to stochastic aspects of the speciation of herbivorous arthropods onto host plants.     

Importance of biotic niches versus drift in a plant‐inhabiting arthropod community depends on rarity and trophic group

Communities are mostly composed of rare species; yet, the factors that determine their patterns of occurrence remain obscure. Theory predicts that, in contrast with common species, the occurrence of rare species will be poorly correlated with environmental variables (niches) and more affected by stochasticity (ecological drift), but how this pattern varies across different trophic groups is still poorly understood. Here, we compared the ability of environmental variables (bottom–up biotic niches) to predict the occurrence of plant‐dwelling arthropods across different abundance classes in the Cape Floristic Region of South Africa. We compared three trophic groups, including 104 herbivorous hemipteran, 171 parasitoid wasp and 84 spider species, totalling 4511 individuals in 48 quadrats. To quantify bottom–up biotic niches, we studied the influences of species composition of plants on hemipterans, and of plants and hemipterans on spiders and wasps. We compared the observed strength of the correlation between rare species and their niches with expectations that were generated by repeatedly rarefying abundant species. A large proportion of arthropod species were very rare, i.e. with only one or two individuals (49–55%). Although rarefying abundant species greatly decreased the correlation with bottom–up biotic niches, bottom–up biotic niches generally better predicted the occurrence of rarefied abundant species than very rare ones, suggesting a greater influence of drift on very rare arthropods. That is, (very) rare arthropods are distributed more randomly than rarefied abundant species. Nevertheless, trophic groups differed in the details of their response to bottom–up biotic niches. Plant species composition was a better predictor of rarefied abundant than truly rare hemipterans. In contrast, the importance of bottom–up biotic niches among abundance classes varied less visibly in spiders and wasps. Our study thus suggests that the importance of niches in structuring arthropod communities depends on species rarity and trophic group.     

Ecological niches of ectoparasites

On mammals and birds communities of ectoparasites are present, which can include scores of ticks, mites and insects species. The parasitizing of arthropods terrestrial vertebrates appeared as far back a the Cretaceous period, and after 70-100 mil. years of the coevolution ectoparasites have assimilated all food resources and localities of the hosts' bodies. To the present only spatial and (to the less extent) trophic niches of parasitic insects, ticks and mites are studied completely enough. The main results these investigations are discussed in the present paper. A high abundance of the communities is reached because of their partition into the number of ecological niches. Host is complex of ecological niches for many ectoparasites species. These niches reiterate in the populations of a species closely related species of hosts and repeat from generation to generation. The each part of host (niche) being assimilated be certain parasite species is available potentially for other species. The partition of host into ecological niches is clearer than the structure of ecosystems including free-living organisms. A real extent of the ecological niches occupation by different species of ticks, mites and insects is considerably lower than a potential maximum. The degree of ecological niches saturation depends on the history of the coevolution of parasites community components, previous colonization be new ectoparasite species and many other ecological factors affecting host-parasite system. The use of the ecological niche conception in parasitology is proved to be rather promising. Ectoparasites communities because of their species diversity, different types of feeding and a number of habitats on host represent convenient models and study of them can contribute significantly to the developmeht of the general conception of ecological niche.     

Effects of cyanogenesis in bracken fern (Pteridium aquilinum) on associated insects

Abstract. 1. The relationship between cyanogenesis in bracken fern and the insect fauna feeding on the plant was investigated over a 3 year period. The most common insects between May and July, while cyanide levels were high, were the sawflies Strongylogaster impressata Provancher, S.multicincta Norton, Aneug-menus fzavipes (Norton), the aphid Macrosiphum euphorbiae (Thomas) and a microlepidopteran species of Monochroa .
2. Collections of insects from cyanogenic and acyanogenic fronds showed significantly fewer sawflies on the cyanogenic fronds. The aphid and the microlepidopteran were randomly distributed with respect to cyanogenicity.
3. Feeding tests for two of the sawfly species ( Simpressata and Smulticincta ) showed that larvae grew more slowly and had a higher mortality when raised on cyanogenic fronds than on acyanogenic ones.
4. Field collected cyanogenic bracken fronds were found to have sustained less damage from chewing herbivores compared with acyanogenic fronds.     

Does the bird cherry have its ‘fair share’ of insect pests? An appraisal of the species—area relationships of the phytophagous insects associated with British Prunus species

Abstract. 1. Field and published data reveal that there are at least twenty-three species of phytophagous insects associated with Prunus padus L. in Britain. Most of these species occur on the tree between May and September. Interspecific competition does not appear to be severe.
2. There are a total of 196 species of phytophagous insects feeding upon the various tissues of P. avium, P. cerasifera, P. cerasus, P. domestica, P. laurocerasus, P. padus and P. spinosa.
3. A highly significant species (5)-distribution (A) relationship of the form In (S+l)=0.92 In A-2.19, was obtained for the Prunus species examined within Britain (excluding Ireland), i.e. the wider the distribution of the plant the greater the number of insect species associated with it.
4. However, critical analysis of the data indicate that P. padus does not support as many different insect species as expected.
5. Neither plant form nor age of establishment within the British Isles accounted for this discrepancy.
6. The possibility of using a plant's insect fauna to describe its relationship to other plants in the same genus is discussed.     

The taxonomy and phytogeography of bracken—a review

Some of the biological problems presented by bracken, Pteridium aquilinum (L.) Kuhn, are posed. Its taxonomic position within the Pteridophyta and the delimitation of entities within the genus are discussed on the basis of morphological and cytological evidence. The geographical ranges of the various brackens world-wide are described and mapped in outline, and emphasis placed on reviewing the natural ecological role of bracken in plant communities throughout the world. Further geographic areas where taxonomic investigation of Pteridium is most needed are indicated, and evidence of the reproductive, dispersal, establishment, colonizing ability and vegetative persistence of bracken is reviewed. Its palaeobiological spread, with associated vegetational history, and the effects on this of anthropogenic influences—better known than are comparable details for any other pteridophyte—are detailed, and the present magnitude of the resulting bracken problem in Britain (and especially in upland Britain) indicated.     

Evolutionary dynamics of specialisation in herbivorous stick insects

Understanding the evolutionary dynamics underlying herbivorous insect mega‐diversity requires investigating the ability of insects to shift and adapt to different host plants. Feeding experiments with nine related stick insect species revealed that insects retain the ability to use ancestral host plants after shifting to novel hosts, with host plant shifts generating fundamental feeding niche expansions. These expansions were, however, not accompanied by expansions of the realised feeding niches, as species on novel hosts are generally ecologically specialised. For shifts from angiosperm to chemically challenging conifer hosts, generalist fundamental feeding niches even evolved jointly with strong host plant specialisation, indicating that host plant specialisation is not driven by constraints imposed by plant chemistry. By coupling analyses of plant chemical compounds, fundamental and ecological feeding niches in multiple insect species, we provide novel insights into the evolutionary dynamics of host range expansion and contraction in herbivorous insects.     

Wolbachia infection frequencies in insects: evidence of a global equilibrium?

Wolbachia are a group of cytoplasmically inherited bacteria that cause reproduction alterations in arthropods, including parthenogenesis, reproductive incompatibility, feminization of genetic males and male killing. Previous general surveys of insects in Panama and Britain found Wolbachia to be common, occurring in 16-22% of species. Here, using similar polymerase chain reaction methods, we report that 19.3% of a sample of temperate North American insects are infected with Wolbachia, a frequency strikingly similar to frequencies found in two other studies in widely separated locales. The results may indicate a widespread equilibrium of Wolbachia infection frequencies in insects whose maintenance remains to be explained. Alternatively, Wolbachia may be increasing in global insect communities. Within each of the three geographic regions surveyed, Hymenoptera are more frequently infected with A group Wolbachia and Lepidoptera more frequently infected with B group Wolbachia.     

Effects of insects on primary production in temperate herbaceous communities: a meta-analysis

Abstract.  1. The effects of insects on primary production in temperate herbaceous communities were investigated in a meta-analysis. The following hypotheses were tested: (1) the effect of insects on primary production depends on community type, (2) the effect of insects on primary production varies as a function of productivity, (3) insects have a greater effect on primary production in communities with low species diversity, and (4) insects have a larger effect on primary production during outbreaks.
2. Data were collected from 24 studies in which insecticides were used to suppress insects in self-sown or pastoral communities. Effect sizes were calculated from sprayed and control plot standing crop or yield, expressed as the log response ratio, ln (sprayed plot phytomass/control plot phytomass).
3. There was a significant increase in primary production as a result of insect suppression. Forb-dominated communities showed a more variable response than graminoid communities. During outbreaks, insects had a greater negative impact on primary production. Effect size was unaffected by productivity or plant species richness.
4. Although insects lower primary production in a diversity of temperate herbaceous communities, the basic measures by which such communities are often described have little effect on the proportional impact that insects have on primary production. While outbreaks are significant predictors of higher negative impact on primary production, causes of outbreaks are not always related to traits of the plant community.     

The structure of the arthropod community on bracken

The species of herbivorous arthropods known to feed on bracken in Britain are listed and the seasonal changes in the structure of this arthropod community are discussed.     

Assessment of the diversity and abundance of terrestrial mangrove arthropods in southern New South Wales,Australia

Abstract The diversity and abundance of arboreal and flying arthropods, in three mangrove patches along the south coast of New South Wales, Australia, was investigated to determine the degree of spatial variability in the assemblages among patches. Intercept traps and restricted canopy fogging were used to sample the communities at Minnamurra, Bonnievale and Kurnell. Twelve orders of arthropods were detected, incorporating 252 morphospecies. Abundance, species richness and species composition were very similar across all patches, the variation being much smaller than expected. These findings suggest that the composition of the arboreal and flying fauna associated with mangrove patches are very similar among patches, but preliminary results also showed that species composition could be highly variable within a patch. Variation between the trapping methods was large, as expected . Intercept trapping and restricted canopy fogging techniques were found to sample different suites of species and therefore complement each other well in sampling programs. Cumulative species curves differed between time periods but generally were flatter for intercept traps than for restricted canopy fogging. Results suggested, for a given level of effort, intercept traps caught a more representative sample of the species composition available to them.     

Microhabitat specialization and similarity of scale-insect assemblages on different fruit trees and in different countries

Abstract. 1. Assemblages of scale-insects on deciduous fruit trees were compared in a wide geographic range of Palaearctic subregions.
2. The parts of all host plant species in the whole geographic region were unsaturated with scale species. There was no clear niche expansion, nor any niche segregation between scale-insect species.
3. These results show that even plants such as deciduous trees, rich in coccoid species, are not fully utilized by scale insects.
4. The similarity of the scale-insect faunas on different plant species could be explained partly by the distribution patterns of the insects, and partly by phylogenetic connection between host plants.     

Arthropod community diversity and trophic structure: a comparison between extremes of plant stress

Abstract.  1. Previous studies have shown that plant stress and plant vigour impact the preference and performance of many insect species. Global climate-change scenarios suggest that some regions such as continental interiors may become increasingly subject to severe drought. In combination, these two issues suggest that drought-driven plant stress may impact insect communities on a landscape scale. While there have been many population studies relating plant stress to the life history of individual herbivore species, far less is known about how plant stress affects entire communities.
2. To study the effect of plant stress on arthropod communities, arthropods were sampled from the canopies of pinyon pines ( Pinus edulis ) growing at sites with a history of chronically high environmental stress (e.g. lower water potentials, soil moisture, and reduced growth rates), and those growing under more favourable conditions. Sampling in these environments yielded >59 000 arthropods, representing 287 species from 14 orders and 80 families, and revealed three major community patterns.
3. First, chronic stress significantly altered community composition. Second, trees growing under high stress supported about 1/10th the number of arthropods, and roughly half the species as trees growing under more favourable conditions. Third, of the 33 abundant herbivore species that exhibited a significantly skewed distribution towards either high- or low-stress trees, 73% were skewed with higher numbers on low-stress trees.
4. The pattern of potentially reduced arthropod diversity and abundance on stressed pines observed in this study may further compound the loss of species resulting from the recent, landscape-scale drought-induced mortality of pines in the southwestern USA.     

Additive and interactive effects of plant genotypic diversity on arthropod communities and plant fitness

Recent research suggests that genetic diversity in plant populations can shape the diversity and abundance of consumer communities. We tested this hypothesis in a field experiment by manipulating patches of Evening Primrose ( Oenothera biennis ) to contain one, four or eight plant genotypes. We then surveyed 92 species of naturally colonizing arthropods. Genetically diverse plant patches had 18% more arthropod species, and a greater abundance of omnivorous and predacious arthropods, but not herbivores, compared with monocultures. The effects of genotypic diversity on arthropod communities were due to a combination of interactive and additive effects among genotypes within genetically diverse patches. Greater genetic diversity also led to a selective feedback, as mean genotype fitness was 27% higher in diverse patches than in monocultures. A comparison between our results and the literature reveals that genetic diversity and species diversity can have similar qualitative and quantitative effects on arthropod communities. Our findings also illustrate the benefit of preserving genetic variation to conserve species diversity and interactions within multitrophic communities.     

Insect diversity in Cape fynbos and neighbouring South African vegetation

Aim  It has often been suggested that South Africa's Cape fynbos shrublands, although extremely rich in plant species, are poor in insects, thus representing a notable exception from the broad plant–insect diversity relationship. The aims of this study were to compare the diversity patterns of plant-inhabiting insects in fynbos and the vegetation of three neighbouring biomes (grassland, subtropical thicket, and Nama-karoo), and to test for a general relationship between plant diversity and insect diversity across these biomes.
Location  South-western to south-eastern South Africa.
Methods  We conducted seasonal plant surveys and sweep insect sampling in 10 × 10 m plots in the Baviaanskloof Conservation Area (Eastern Cape), where all four biomes occur. We also conducted once-only collections in the core area of each biome.
Results  Fynbos plots had insect diversity values similar to those of grassland and subtropical thicket (a dense, evergreen and spinescent shrubland with a high abundance of succulents and climbers), and significantly higher than Nama-karoo (an open, semiarid shrubland). A remarkably strong positive relationship was found between plant and insect species richness.
Main conclusions  Previous generalizations were based on a few insect groups (e.g. butterflies, under-represented in fynbos), but ignored published results on other groups (e.g. galling insects, which are in fact over-represented in this vegetation). We show that, overall, insect diversity in fynbos is comparable to that of neighbouring biomes. Fynbos vegetation does not represent a significant exception from the broad positive relationship between plant diversity and insect diversity.     

Predicting patterns of plant species richness in megadiverse South Africa

Using new tools (boosted regression trees) in predictive biogeography, with extensive spatial 23 distribution data for >19 000 species, we developed predictive models for South African plant species richness patterns. Further, biome level analysis explored possible functional determinants of country‐wide regional species richness. Finally, to test model reliability independently, we predicted potential alien invasive plant species richness with an independent dataset. Amongst the different hypotheses generally invoked to explain species 30 diversity (energy, favorableness, topographic heterogeneity, irregularity and seasonality), results revealed topographic heterogeneity as the most powerful single explanatory variable for indigenous South African plant species richness. Some biome‐specific responses were observed, i.e. two of the five analyzed biomes (Fynbos and Grassland) had richness best explained by the “species‐favorableness” hypothesis, but even in this case, topographic heterogeneity was also a primary predictor. This analysis, the largest conducted on an almost exhaustive species sample in a species‐rich region, demonstrates the preeminence of topographic heterogeneity in shaping the spatial pattern of regional plant species richness. Model reliability was confirmed by the considerable predictive power for alien invasive species richness. It thus appears that topographic heterogeneity controls species richness in two main ways: firstly, by providing an abundance of ecological niches in contemporary space (revealed by alien invasive species richness relationships) and secondly, by facilitating the persistence of ecological niches through time. The extraordinary richness of the South African Fynbos biome, a world‐renowned hotspot of biodiversity with the steepest environmental gradients in South Africa, may thus have arisen through both mechanisms. Comparisons with similar regions of the world outside South Africa are needed to confirm the generality of topographic heterogeneity and favorableness as predictors of plant richness.     

Phytophagous British insects and mites and their food-plant families: total numbers and polyphagy

The Phytophagous Insects Data Base (PIDB) was used to summarize information about 6933 species of British insects/mites and their food-plant families. Total species in 183 insect families on 127 vascular plant families were correlated with numbers of plant species in each family in Britain and Europe, accounting for 41.8% of the variation. Families with trees had more insects, while ferns and aquatic, uncommon and ephemeral plants, particularly orchids, had fewer. Seven individual insect families were analysed separately: Aphididae were more closely correlated with plant species numbers than were the more polyphagous Geometridae. Agromyzidae were uncommon on families with trees. Of insect species 75.8% fed on only one plant family and 10.1% were recorded on two families. Species feeding on three families or more were considered to be polyphagous. In the 30 largest insect families, eight had species feeding on only one plant family, while 12 had more than 20% of their species polyphagous. Proportions of family-restricted insects were high on Pinaceae, Gramineae and Compositae, but low on Corylaceae, Primulaceae and Tiliaceae. More pest species were polyphagous. Polyphagy was related to large larvae, larvae overwintering, wood-feeding, ephemeral food sources, poor dispersal abilities and omnivory. Some polyphagous species might be actively evolving in exploitation of vacant niches, leading to later progressive specialization to food-plants.     

我国烟田昆虫群落的划分及其特征分析

根据纬度、海拔、温度、降雨、日照等地理、气象因子指标,对我国71个植烟市和(或)县及生态条件独特的市和(或)县进行了聚类分析,聚类结果与我国烟草的7个一级种植区基本吻合,我国烟田烟草昆虫群落可依此划分为东北、黄淮、长江中上游、长江中下游、西南、南部和北部西部等7个。各群落的重要烟草昆虫种类有所不同,烟草昆虫天敌的种类差别更大。其中西南群落食物网最复杂。     

Herbivore-induced responses and patch heterogeneity affect abundance of arthropods on plants

Abstract.  1. Plants respond to herbivore damage by inducing defences that can affect the abundance of herbivores and predators. These tritrophic interactions may be influenced by heterogeneity in plant neighbourhood.
2. In the present study, the effects of induced responses on the abundance of herbivores (flea beetles and aphids), omnivores (pirate bugs and thrips), and predators (lady beetles and spiders) on individual plants and their neighbours between and within patches composed of three tomato plants was investigated.
3. Herbivore damage was manipulated to create homogeneous patches where either all or none of the plants had defences induced by herbivore damage, and heterogeneous patches where only one of the plants was induced.
4. Arthropod abundance on plants at different scales was compared by testing between patch effects (patch level), for neighbourhood effects at the plant phenotype level (neighbourhood level), and between near and far plants (within patch position).
5. At the patch level , plants in homogeneously induced patches contained fewer flea beetles and pirate bugs, but more lady beetles, compared with homogeneously non-induced patches. There was no effect of patch type on the abundance of aphids, thrips, and spiders on plants.
6. At the neighbourhood level , induced plants in heterogeneous patches contained more flea beetles and pirate bugs compared with induced plants in homogeneous patches, indicating that the abundance of some herbivores and omnivores on induced plants varied depending on the phenotype of the other plants within the patch. Within patch position, there was no evidence that the abundance of herbivores or predators on non-induced plants was affected by proximity to an induced plant.
7. Therefore, variation in plant neighbourhood generated by induced plant responses affected the abundance of three arthropods from three feeding guilds.