Arthropods associated with fungal galls: do large galls support more abundant and diverse inhabitants?

Fungus-induced galls can attract spore-feeding arthropods as well as gall-feeding ones, resulting in diverse communities. Do large fungal galls support more abundant and diverse arthropod communities than small fungal galls? To address this question, we investigated the structure of the arthropod community associated with bud galls induced by the fungus Melanopsichium onumae on the tree species Cinnamomum yabunikkei (Lauraceae) in central Japan. Thirteen species of arthropods were associated with M. onumae galls. Dominant arthropod species were represented by the larvae of a salpingid beetle (a spore feeder), a nitidulid beetle (a spore feeder), a cosmopterigid moth (a spore feeder), an unidentified moth (a gall tissue feeder), and a drosophilid species (a gall tissue feeder). Arthropod abundance and species richness were positively correlated with gall diameter. The majority of the most abundant species were more frequently found in large galls than in small ones, indicating that large fungal galls, which have more food and/or space for arthropods, could support a more abundant and diverse arthropod community.     

Effects of environmental parameters on the chestnut gall wasp and its complex of indigenous parasitoids

The chestnut gall wasp (CGW), Dryocosmus kuriphilus, an invasive pest native to China, has caused severe yield and economic losses to chestnut production in Europe since its arrival in 2002. In Southern Italy, the complex of indigenous parasitoids colonizing CGW was monitored between 2013 and 2015, with the aim of estimating the composition of the indigenous parasitoid complex, its ability to control CGW populations, and the interactions of both factors with several measured environmental parameters. We compared results among three differently managed field types. Results showed an increase in the rate of parasitism both when the host population density was lower and in unmanaged chestnut stands with more natural conditions. The percentage of parasitism in galls was related to morphological traits of the galls and to higher seasonal temperatures, which reduced the parasitism intensity because CGW develops earlier under such conditions. The host–parasitoid mortality inside galls varied among sites and was associated mostly with rot fungi during wet spring and summer months. Parasitoid species richness was similar among the study sites, but the proportion of parasitoid species differed between orchards and unmanaged coppice stands. The timing of attack by parasitoids followed a species-specific successional sequence throughout the larva-to-adult life cycle of the CGW. These interactions should be considered in future research on trophic relationships and when modeling invasive scenarios for new pest species.     

Host manipulation by the orange leafhopper Cicadulina bipunctata: gall induction on distant leaves by dose-dependent stimulation

The evolution of the gall-inducing ability in insects and the adaptive significance of the galling habit have been addressed by many studies. Cicadulina bipunctata, the maize orange leafhopper, is an ideal study organism for evaluating these topics because it can be mass-reared and it feeds on model plants such as rice (Oryza sativa) and maize (Zea mays). To reveal differences between gall inductions by C. bipunctata and other gall inducers, we conducted four experiments concerning (a) the relationship between the feeding site and gall-induction sites of C. bipunctata on maize, (b) the effects of leafhopper sex and density, (c) the effects of length of infestation on gall induction, and (d) the effects of continuous infestation. C. bipunctata did not induce galls on the leaves where it fed but induced galls on other leaves situated at more distal positions. The degree of gall induction was significantly correlated with infestation density and length. These results indicate that C. bipunctata induces galls in a dose-dependent manner on leaves distant from feeding sites, probably by injecting chemical(s) to the plant during feeding. We suggest that insect galls are induced by a chemical stimulus injected by gall inducers during feeding into the hosts.     

Colour mimicry and sexual deception by Tongue orchids (Cryptostylis)

Typically, floral colour attracts pollinators by advertising rewards such as nectar, but how does colour function when pollinators are deceived, unrewarded, and may even suffer fitness costs? Sexually deceptive orchids are pollinated only by male insects fooled into mating with orchid flowers and inadvertently transferring orchid pollinia. Over long distances, sexually deceptive orchids lure pollinators with counterfeit insect sex pheromones, but close-range deception with colour mimicry is a tantalising possibility. Here, for the first time, we analyse the colours of four sexually deceptive Cryptostylis orchid species and the female wasp they mimic (Lissopimpla excelsa, Ichneumonidae), from the perspective of the orchids’ single, shared pollinator, male Lissopimpla excelsa. Despite appearing different to humans, the colours of the orchids and female wasps were effectively identical when mapped into a hymenopteran hexagonal colour space. The orchids and wasps reflected predominantly red-orange wavelengths, but UV was also reflected by raised bumps on two orchid species and by female wasp wings. The orchids’ bright yellow pollinia contrasted significantly with their overall red colour. Orchid deception may therefore involve accurate and species-specific mimicry of wavelengths reflected by female wasps, and potentially, exploitation of insects’ innate attraction to UV and yellow wavelengths. In general, mimicry may be facilitated by exploiting visual vulnerabilities and evolve more readily at the peripheries of sensory perception. Many sexually deceptive orchids are predominantly red, green or white: colours that are all potentially difficult for hymenoptera to detect or distinguish from the background.     

Host pollination mode and mutualist pollinator presence: net effect of internally ovipositing parasite in the fig–wasp mutualism

The Ficus–their specific pollinating fig wasps (Chalcidoidea, Agaonidae) interaction presents a striking example of mutualism. Figs also shelter numerous non-pollinating fig wasps (NPFW) that exploit the fig–pollinator mutualism. Only a few NPFW species can enter figs to oviposit, they do not belong to the pollinating lineage Agaonidae. The internally ovipositing non-agaonid fig wasps can efficiently pollinate the Ficus species that were passively pollinated. However, there is no study to focus on the net effect of these internally ovipositing non-agaonid wasps in actively pollinated Ficus species. By collecting the data of fig wasp community and conducting controlled experiments, our results showed that internally ovipositing Diaziella bizarrea cannot effectively pollinate Ficus glaberrima, an actively pollinated monoecious fig tree. Furthermore, D. bizarrea failed to reproduce if they were introduced into figs without Eupristina sp., the regular pollinator, as all the figs aborted. Furthermore, although D. bizarrea had no effect on seed production in shared figs, it significantly reduced the number of Eupristina sp. progeny emerging from them. Thus, our experimental evidence shows that reproduction in Diaziella depends on the presence of agaonid pollinators, and whether internally ovipositing parasites can act as pollinators depends on the host fig’s pollination mode (active or passive). Overall, this study and others suggest a relatively limited mutualistic role for internally ovipositing fig wasps from non-pollinator (non-Agaonidae) lineages.     

Heat-balling wasps by honeybees

Defensiveness of honeybee colonies of Apis cerana and Apis mellifera (actively balling the wasps but reduction of foraging) against predatory wasps, Vespa velutina, and false wasps was assessed. There were significantly more worker bees in balls of the former than latter. Core temperatures in a ball around a live wasp of A. cerana were significantly higher than those of A. mellifera, and also significantly more when exposed to false wasps. Core temperatures of bee balls exposed to false wasps were significantly lower than those exposed to V. velutina for both A. cerana and for A. mellifera. The lethal thermal limits for V. velutina, A. cerana and A. mellifera were significantly different, so that both species of honeybees have a thermal safety factor in heat-killing such wasp predators. During wasps attacks at the hives measured at 3, 6 and 12 min, the numbers of Apis cerana cerana and Apis cerana indica bees continuing to forage were significantly reduced with increased wasp attack time. Tropical lowland A. c. indica reduced foraging rates significantly more than the highland A. c. cerana bees; but, there was no significant effect on foraging by A. mellifera. The latency to recovery of honeybee foraging was significantly greater the longer the duration of wasp attacks. The results show remarkable thermal fine-tuning in a co-evolving predator–prey relationship.     

Overstorey tree density and understorey regrowth effects on plant composition,stand structure and floristic richness in grazed temperate woodlands in eastern Australia

As natural woodlands decline in both extent and quality worldwide, there is an increasing recognition of the biodiversity conservation value of production landscapes. In low-input, low-productivity grazing systems in Australia, the modification of natural woodlands through overstorey tree and woody regrowth removal are vegetation management options used by landholders to increase native grass production for livestock grazing; however, there is little empirical evidence to indicate at what tree densities biodiversity attributes are compromised. We examined the effects of overstorey tree density and understorey regowth on the floristic composition, stand structure and species richness of eucalypt woodlands in a grazing landscape in the Traprock region of southern Queensland, Australia. We sampled 47 sites stratified according to vegetation type (Eucalyptus crebra/Eucalyptus dealbata woodland; Eucalyptus melliodora/Eucalyptus microcarpa grassy woodland), density of mature trees (<6 trees/ha; 6–20 trees/ha; >20 trees/ha), and presence/absence of regrowth. Distinct patterns in composition were detected using indicator species analysis and non-metric multidimensional scaling, with low density areas compositionally indistinguishable, although distinct from other land management units. Within vegetation type, medium tree density woodlands were compositionally similar to high density and reference woodlands. Species richness ranged from 18 to 67 species per 500 m² across all sites. No differences in total or native species richness were detected across management units; however, some differences in exotic species richness were detected. Differences in grass cover existed between low and high density management units, yet no difference in grass cover was evident between low and medium density management units. Our results suggest that medium tree densities may provide biodiversity benefits concordant with more natural areas, yet not adversely impact on pasture production. Retaining trees in grazing landscapes provides significant landscape heterogeneity and important refuges for species that may be largely excluded from open grassland habitats. Maintaining a medium density of overstorey trees in grazed paddocks can provide both production and biodiversity benefits.     

Hydrocarbons from plants: Analytical methods and observations

We have suggested that certain plants rich in hydrocarbon-like materials might be cultivated for renewable photosynthetic products. Two species were selected for experimental plantations: Euphorbia lathyris, an annual from seed and Euphorbia tirucalli, a perennial from cuttings. The yield from each species is over 10 barrels of oil/acre/year without genetic or agronomic improvement. In addition to plants, there are trees, such as species of Copaifera in Brazil and other tropical areas, which produce a diesel-like oil upon tapping. Each tree produces approximately 40 liters of hydrocarbon per year, and this material can be used directly by a diesel-powered car. Further efforts to develop plants as alternate energy sources are underway, as well as a continuing search for additional plant species throughout the world which have a similar capability.     

Different patterns of oviposition learning in two closely related ectoparasitoid wasps with contrasting reproductive strategies

Many parasitoid wasps learn host-associated cues and use them in subsequent host-searching behavior. This associative learning, namely “oviposition learning,” has been investigated in many studies. However, few studies have compared multiple species, and no comparative study has previously been conducted on ectoparasitoid species. We compared the effects of oviposition learning on host preference and offspring sex ratio in two closely related ectoparasitoid wasps with contrasting reproductive strategies, Anisopteromalus calandrae (r-strategist) and its sibling species (K-strategist). Using two bruchine hosts, Callosobruchus chinensis and Callosobruchus maculatus larvae infesting the cowpea Vigna unguiculata, oviposition choice experiments were performed at high and low host densities. In both species, no conspicuous effect on the offspring sex ratio was detected, but effects on host preference were found to differ between the species. In A. calandrae, the effects were detected only at high host density, suggesting that oviposition learning plays a role in host discrimination from a short distance but not from a long distance. In the sibling species, those effects were not detected in any of the cases, suggesting the absence of oviposition learning. These results are compatible with those of previous comparative studies of endoparasitoid wasps in that few lifetime oviposition experiences and/or low reward per foraging decision result in low or absent oviposition learning ability. This finding may indicate that ecological traits contributing to learning ability are similar between endoparasitoid and ectoparasitoid wasps. Thus, our species comparison of ectoparasitoids provides another model system for investigating learning and memory dynamics in parasitoid wasps.     

Exploring public perceptions of solutions to tree diseases in the UK: Implications for policy-makers

Tree diseases are on the increase in many countries and the implications of their appearance can be political, as well as ecological and economic. Preventative policy approaches to tree diseases are difficult to formulate because dispersal pathways for pest and pathogens are numerous, poorly known and likely to be beyond human management control. Genomic techniques could offer the quickest and most predictable approach to developing a disease tolerant native ash.The population of European Ash (Fraxinus Excelsior) has suffered major losses in the last decade, due to the onset of Hymenoscyphus fraxineus (previously called Chalara Fraxinea) commonly known in the UK as ash dieback. This study presents evidence on the public acceptability of tree-breed solutions to the spread of Chalara, with the main aim to provide science and policy with an up-stream ‘steer’ on the likely public acceptability of different tree breeding solutions. The findings showed that whilst there was a firm anti-GM and ‘we shouldn’t tamper with nature’ attitude among UK publics, there was an equally firm and perhaps slightly larger pragmatic attitude that GM (science and technology) should be used if there is a good reason to do so, for example if it can help protect trees from disease and help feed the world. The latter view was significantly stronger among younger age groups (Millennials), those living in urban areas and when the (GM)modified trees were destined for urban and plantation, rather than countryside settings. Overall, our findings suggest that the UK government could consider genomic solutions to tree breeding with more confidence in the future, as large and influential publics appear to be relaxed about the use of genomic techniques to increase tolerance of trees to disease.     

The hunter becomes the hunted: when cleptobiotic insects are captured by their target ants

Here we show that trying to rob prey (cleptobiosis) from a highly specialized predatory ant species is risky. To capture prey, Allomerus decemarticulatus workers build gallery-shaped traps on the stems of their associated myrmecophyte, Hirtella physophora. We wondered whether the frequent presence of immobilized prey on the trap attracted flying cleptoparasites. Nine social wasp species nest in the H. physophora foliage; of the six species studied, only Angiopolybia pallens rob prey from Allomerus colonies. For those H. physophora not sheltering wasps, we noted cleptobiosis by stingless bees (Trigona), social wasps (A. pallens and five Agelaia species), assassin bugs (Reduviidae), and flies. A relationship between the size of the robbers and their rate of capture by ambushing Allomerus workers was established for social wasps; small wasps were easily captured, while the largest never were. Reduviids, which are slow to extract their rostrum from prey, were always captured, while Trigona and flies often escaped. The balance sheet for the ants was positive vis-à-vis the reduviids and four out of the six social wasp species. For the latter, wasps began by cutting up parts of the prey’s abdomen and were captured (or abandoned the prey) before the entire abdomen was retrieved so that the total weight of the captured wasps exceeded that of the prey abdomens. For A. pallens, we show that the number of individuals captured during attempts at cleptobiosis increases with the size of the Allomerus’ prey.     

Geographic variations in seed dispersal by ants: are plant and seed traits decisive?

The effect of local ant species on the dispersal success of a myrmecochorous plant, Helleborus foetidus, was analyzed in two populations of the Iberian Peninsula (Caurel and Cazorla, respectively). The contribution of the various local ant species to dispersal was very unequal. While 5 and 19 ant taxa visited the plants of Caurel and Cazorla, respectively, most removal activity (67 and 80%) was performed by two species only (Formica lugubris and Camponotus cruentatus, respectively). Visits by dispersers were also unequally distributed between neighboring plants. While some plants were always visited during the period of seed release, others were never visited. A regression model indicated that this pattern might be explained by two plant traits: ants preferred to visit plants that released more seeds and whose elaiosomes were richer in oleic acid. Although it has long been known that this compound triggers removal by ants, it is the first demonstration that quantitative variations in elaiosome traits contribute to variation in dispersal success. Finally, other variables being equal, morphological traits (seed size, elaiosome size, and elaiosome/seed size ratio) did not affect ant behavior. Although myrmecochory has long been considered a diffuse interaction, our results support the idea that, at local scale, a limited number of ant species may be decisive to its evolution.     

Tracking parasitoids with the stable isotope 44Ca in agroecosystems

Habitat management influences spatial and temporal distribution of parasitoids in farmland. The current work evaluates, for the first time, the potential of a novel marking technique using a calcium stable isotope (⁴⁴Ca) under field conditions. In two subsequent trials, ⁴⁴Ca-enriched Cotesia glomerata parasitoids were released into an organically managed cabbage field in a region known to harbor natural populations of this species. The trap plants infested with Pieris brassicae host larvae were distributed in the trial areas and collected 3 days after parasitoid release. Parasitism by released wasps was determined through calcium isotope analysis of the recovered caterpillars using inductively coupled plasma mass spectrometry (ICPMS). The spatial habitat use by female parasitoids was determined based on marked caterpillars on the trap plants. Both trials yielded relatively consistent results, showing that C. glomerata females dispersed over at least 50 m within 3 days. The total proportion of caterpillars parasitized by the marked wasps amounted to 32.4 and 24.4%, respectively, with no statistically significant difference. The potential of this approach for field investigations on habitat management and biological control is discussed.     

Effects of road age on the structure of roadside vegetation in south-eastern Australia

In many agricultural landscapes, roadside (or road verge) environments provide important refuge for threatened native species and ecosystems, and are often selected as benchmark sites to guide restoration activities. However few studies have investigated potential temporal variability in roadside vegetation conditions. In this paper we used archived cadastral maps to determine road age and examine potential variability in roadside vegetation structures in a typical rural landscape in south-eastern Australia. We found significant differences in the density of mature trees for road segments in different road-age categories. The oldest roads (<1870s) were characterized by having the greatest density of large hollow-bearing Eucalyptus trees, but few native conifer trees or shrubs. Roads surveyed when broad-scale clearing commenced (1870–1879), and not the oldest roads, were found to be more intact in terms of the density of large pre-settlement trees, range of tree stem-size classes and overall shrub diversity. By contrast, the youngest roads (post-1900s) had the greatest number of native conifer trees, but few shrubs or large Eucalyptus trees. As a result, roads of different ages had different densities of hollow-bearing trees, which is discussed in terms of past land-use legacies. These results have important implications for selecting roadsides as benchmark sites for restoration activities, and highlight the critical importance of roadsides to conserving native biota in agricultural landscapes.     

Bee-hawking by the wasp, <Emphasis Type="BoldItalic">Vespa velutina</Emphasis>, on the honeybees <Emphasis Type="BoldItalic">Apis cerana</Emphasis> and <Emphasis Type="BoldItalic">A</Emphasis>. <Emphasis Type="BoldItalic">mellifera</Emphasis>

The vespine wasps, Vespa velutina, specialise in hawking honeybee foragers returning to their nests. We studied their behaviour in China using native Apis cerana and introduced A. mellifera colonies. When the wasps are hawking, A. cerana recruits threefold more guard bees to stave off predation than A. mellifera. The former also utilises wing shimmering as a visual pattern disruption mechanism, which is not shown by A. mellifera. A. cerana foragers halve the time of normal flight needed to dart into the nest entrance, while A. mellifera actually slows down in sashaying flight manoeuvres. V. velutina preferentially hawks A. mellifera foragers when both A. mellifera and A. cerana occur in the same apiary. The pace of wasp-hawking was highest in mid-summer but the frequency of hawking wasps was three times higher at A. mellifera colonies than at the A. cerana colonies. The wasps were taking A. mellifera foragers at a frequency eightfold greater than A. cerana foragers. The final hawking success rates of the wasps were about three times higher for A. mellifera foragers than for A. cerana. The relative success of native A. cerana over European A. mellifera in thwarting predation by the wasp V. velutina is interpreted as the result of co-evolution between the Asian wasp and honeybee, respectively.     

Reproductive division of labour coevolves with gall size in Australian thrips with soldiers

An analysis of multiple species of Australian gall-inducing thrips with soldiers reveals a significant negative correlation between the size of gall produced and the reproductive division of labour. This correlation suggests that the evolution of smaller galls limited the available space and feeding sites for the offspring of female soldiers, and was a major factor that led to the evolution of an altruistic caste in the gall-inducers. We argue that high levels of inbreeding by singly mated foundresses and incestuous mating by her soldier offspring are key to this evolutionary relationship because they make the relatedness of a female soldier to her daughters and sisters approximately equal. Evidence that relatedness plays an important role is strengthened by the observation of outbred multiply mated foundresses and unbiased sex ratio of dispersers in Oncothrips waterhousei, and the inference that both gall volume and skew decreased along this lineage.     

The population ecology of Datura ferox in soybean crops. A simulation approach incorporating seed dispersal

A numerical model for simulating the population dynamics of Datura ferox L. (chamico, chinese thronapple) has been built based on previously reported data. In the model, a soybean field is divided into 0.7×0.7-m conceptual modules. A seed production sub-model simulates the annual seed output of each module, and a seed dispersal sub-model simulates the distribution of these propagules within the field in accordance to a specified dispersal pattern. Different model scenarios were generated by varying the proportion of seeds lost from the soil bank, the annual recruitment, the seedling mortality and the proportion of seeds exported from the field during crop harvest. The results obtained by simulation suggest that (1) seed dispersal due to crop harvesting tends to produce an exponential growth of weed seed production, (2) limited or no success could be attained in the control of D. ferox using procedures that kill the seedlings efficiently if combine harvesters are not adjusted so as to maximize the proportion of weed seeds that are exported from the field and (3) if cleaning debris continues to be returned to the ground during crop harvesting, the improvement in the efficiency of the grain/weed separating mechanisms does not provide an effective long-term strategy to avoid grain contamination problems.     

Shade tree management affects fruit abortion, insect pests and pathogens of cacao

The mortality of cacao fruits caused by early fruit abortion or insect and pathogen attacks was investigated in differently managed agroforestry systems in Central Sulawesi, Indonesia. Nine agroforestry systems shaded by three different types of tree stands were selected, which represented a decrease in structural heterogeneity: forest remnants, diverse planted trees and one or two species of planted leguminose trees. After standardized manual cross-pollination, the development of 600 fruits on 54 trees (6 trees per agroforest) was followed during 18 weeks of fruit development. In total, 432 of all fruits were lost before maturity, which seriously undermined yields. The proportion of harvested fruits per tree (overall average: 27 ± 4%) was not affected by canopy type. Although shade cover did not have a significant effect, losses due to fruit abortion were most likely under forest shade, where nitrogen-fixing leguminose shade trees were absent. Fruit losses due to pathogenic infections and insect attacks increased with the homogenization of the agroforests, supporting the hypothesis that agricultural homogenization increases risks of pest outbreaks. In conclusion, shade management may be improved to increase yields from cacao using highly diversified natural shade agroforestry systems.     

模拟酸雨对亚热带三个树种凋落叶分解速率及分解酶活性的影响

利用凋落物袋法研究了模拟酸雨对中国亚热带地区3个典型树种杉木(Cunninghamia lanceolata)、香樟(Cinnamomum camphora)、银杏(Ginkgo biloba)凋落叶分解速率及酶活性的影响.酸雨分别设置为重度酸雨(pH=2.5)、中度酸雨(pH=4.0)、对照处理(pH =5.6)3个梯度.试验结果表明:酸雨胁迫会抑制凋落叶的分解,且随着酸雨强度的增强,抑制作用更明显.杉木、香樟、银杏3个树种对照处理下周转期分别比中度酸雨短33％、43％、14％,比重度酸雨短44％、52％、17％.3种分解酶对酸雨胁迫的表现不同,脲酶和纤维素酶表现为一定的抑制作用,基本呈现为对照处理(pH=5.6)＞中度酸雨(pH=4.0)＞重度酸雨(pH=2.5),与凋落物的分解速率排列情况一致,蔗糖酶在酸雨胁迫下反而有激活的趋势,表现为重度酸雨(pH=2.5)＞中度酸雨(pH =4.0)＞对照处理(pH=5.6).不同树种对酸雨胁迫的影响有一定的差异性,针叶树种杉木受影响最大,阔叶树种香樟和银杏受影响较小,银杏的抗酸性较强,且针叶树种的分解速率比阔叶树种慢.脲酶和纤维素酶对凋落叶的分解贡献较大,而蔗糖酶的影响相对较小.酶活性与季节有较大的关系,夏季活性相对偏高,冬季活性相对偏低.     

‘Special agents’ trigger social waves in giant honeybees (Apis dorsata)

Giant honeybees (Apis dorsata) nest in the open and have therefore evolved a variety of defence strategies. Against predatory wasps, they produce highly coordinated Mexican wavelike cascades termed ‘shimmering’, whereby hundreds of bees flip their abdomens upwards. Although it is well known that shimmering commences at distinct spots on the nest surface, it is still unclear how shimmering is generated. In this study, colonies were exposed to living tethered wasps that were moved in front of the experimental nest. Temporal and spatial patterns of shimmering were investigated in and after the presence of the wasp. The numbers and locations of bees that participated in the shimmering were assessed, and those bees that triggered the waves were identified. The findings reveal that the position of identified trigger cohorts did not reflect the experimental path of the tethered wasp. Instead, the trigger centres were primarily arranged in the close periphery of the mouth zone of the nest, around those parts where the main locomotory activity occurs. This favours the ‘special-agents’ hypothesis that suggest that groups of specialized bees initiate the shimmering.