Behavioural function and development of body-to-limb proportions and active movement ranges in three stick insect species.

Overall body proportions and relative limb length are highly characteristic for most insect taxa. In case of the legs, limb length has mostly been discussed with regard to parameters of locomotor performance and, in particular cases, as an adaptation to environmental factors or to the mating system. Here, we compare three species of stick and leaf insects (Phasmatodea) that differ strongly in the length ratio between antennae and walking legs, with the antennae of Medauroidea extradentata being much shorter than its legs, nearly equal length of antennae and legs in Carausius morosus, and considerably longer antennae than front legs in Aretaon asperrimus. We show that that relative limb length is directly related to the near-range exploration effort, with complementary function of the antennae and front legs irrespective of their length ratio. Assuming that these inter-species differences hold for both sexes and all developmental stages, we further explore how relative limb length differs between sexes and how it changes throughout postembryonic development. We show that the pattern of limb-to-body proportions is species-characteristic despite sexual dimorphism, and find that the change in sexual dimorphism is strongest during the last two moults. Finally, we show that antennal growth rate is consistently higher than that of front legs, but differs categorically between the species investigated. Whereas antennal growth rate is constant in Carausius, the antennae grow exponentially in Medauroidea and with a sudden boost during the last moult in Aretaon.

Two new species of Oxyethira Eaton 1873 (Trichoptera: Hydroptilidae) from Roraima state, Brazil.

Two new species of Oxyethira from Brazil are described: Oxyethira (Dampfitrichia) jiquitaia sp. nov., and Oxyethira (Tanytrichia) librina sp. nov. Illustrations of male genitalia are provided with each description. These results increase the number of Oxyethira species in Brazil from 37 to 39.

Increasing temperatures reduce invertebrate abundance and slow decomposition.

Decomposition is an essential ecosystem service driven by interacting biotic and abiotic factors. Increasing temperatures due to climate change can affect soil moisture, soil fauna, and subsequently, decomposition. Understanding how projected climate change scenarios will affect decomposition is of vital importance for predicting nutrient cycling and ecosystem health. In this study, we experimentally addressed the question of how the early stages of decomposition would vary along a gradient of projected climate change scenarios. Given the importance of biodiversity for ecosystem service provisioning, we measured the effect of invertebrate exclusion on red maple (Acer rubrum) leaf litter breakdown along a temperature gradient using litterbags in warming chambers over a period of five weeks. Leaf litter decomposed more slowly in the warmer chambers and in the litterbag treatment that minimized invertebrate access. Moreover, increasing air temperature reduced invertebrate abundance and richness, and altered the community composition, independent of exclusion treatment. Using structural equation models, we were able to disentangle the effects of average air temperature on leaf litter loss, finding a direct negative effect of warming on the early stages of decomposition, independent of invertebrate abundance. This result indicates that not only can climate change affect the invertebrate community, but may also directly influence how the remaining organisms interact with their environment and their effectiveness at provisioning ecosystem services. Overall, our study highlights the role of biodiversity in maintaining ecosystem services and contributes to our understanding of how climate change could disrupt nutrient cycling.

Chemical compositions and repellent activity of Clerodendrum bungei Steud. essential oil against three stored product insects.

BACKGROUND: Several species of Verbenaceae have been widely used in medicine, and some species of Verbenaceae have been observed good insecticidal activity, such as Lantana camara and Vitex negundo. There is no report about repellent activity of Clerodendrum bungei Steud. (C. bungei) against stored product insects. The chemical composition of C. bungei essential oil (EO) were identified, repellent activity of methanol extract, EO of C. bungei and two main components of EO against T. castaneum, L. serricorne and L. bostrychophila were evaluated for the first time. RESULTS: EO of C. bungei was obtained by hydrodistillation and analyzed by gas chromatography-mass spectrometry (GC-MS) and GC. A total of 25 components of the C. bungei EO were identified. The principal compounds in the EO were myristicin (75.0%), 2,2,7,7-Tetramethyltricyclo[6.2.1.0(1,6)]undec-4-en-3-one (4.1%) and linalool (3.4%). Results of bioassays indicated that C. bungei EO exerted strong repellent activity against three target insects. As main constituents, myristicin and linalool also had certain repellency. CONCLUSION: This work suggests that the EO of C. bungei has promising potential to develop into botanical repellents for the control of pest damage in warehouses and grain stores.

emTimnewia/em García Aldrete (Psocodea: 'Psocoptera': Ptiloneuridae): new species and variation in the wing venation of emT. jeaneae/em Silva-Neto, García Aldrete amp; Rafael.

A new species of Timnewia from the Brazilian state of Amazonas, is here described, illustrated and named as: Timnewia amazonense n. sp. This new species differs from T. greeni and T. jeaneae in details of the hypandrium and phallosome. New observations on variation in the forewing venation of T. jeaneae are also presented.

Penetration mechanics of elongated female and male genitalia of earwigs.

We unveiled the penile penetration mechanics of two earwig species, Echinosoma horridum, whose intromittent organ, termed virga, is extraordinarily long, and E. denticulatum, whose virga is conversely short. We characterised configuration, geometry, material and bending stiffness for both virga and spermatheca. The short virga of E. denticulatum has a material gradient with the stiffer base, whereas the long virga of E. horridum and the spermathecae of both species are homogeneously sclerotised. The long virga of E. horridum has a lower bending stiffness than the spermatheca. The virga of E. denticulatum is overall less flexible than the spermatheca. We compared our results to a previous study on the penetration mechanics of elongated beetle genitalia. Based on the comparison, we hypothesised that the lower stiffness of the male intromittent organ comparing to the corresponding female structure is a universal prerequisite for the penetration mechanics of the elongated intromittent organ in insects.

A Stresipteran parasite extends the lifespan of workers in a social wasp.

In social wasps, female lifespan depends on caste and colony tasks: workers usually live a few weeks while queens as long as 1 year. Polistes dominula paper wasps infected by the strepsipteran parasite Xenos vesparum avoid all colony tasks, cluster on vegetation where parasite dispersal and mating occur, hibernate and infect the next generation of wasp larvae. Here, we compared the survival rate of infected and uninfected wasp workers. Workers' survival was significantly affected by parasite sex: two-third of workers parasitized by a X. vesparum female survived and overwintered like future queens did, while all workers infected by a X. vesparum male died during the summer, like uninfected workers that we used as controls. We measured a set of host and parasite traits possibly associated with the observed lifespan extension. Infected overwintering workers had larger fat bodies than infected workers that died in the summer, but they had similar body size and ovary development. Furthermore, we recorded a positive correlation between parasite and host body sizes. We hypothesize that the manipulation of worker's longevity operated by X. vesparum enhances parasite's fitness: if workers infected by a female overwinter, they can spread infective parasite larvae in the spring like parasitized gynes do, thus contributing to parasite transmission.

The dynamic eggs of the Phasmatodea and their apparent convergence with plants.

The egg stages of animal life cycles are underappreciated in terms of their capacity for dispersal, protection, and biotic and abiotic interactions. Some of the most intriguing egg morphologies are seen in stick and leaf insects (Phasmatodea). Phasmids are charismatic insects, particularly due to their incredible camouflage, though a lesser-known fact is that their eggs are incredibly diverse in shape and structure, reflecting varying ecological niches. Perhaps most remarkable are those eggs which appear to resemble plant seeds in both their appearance and means of dispersal, such as via water and animal vectors. Numerous hypotheses surrounding the function of these egg morphologies and their apparent convergence with seeds have been proposed; however, empirical evidence remains lacking. Here, we present an initial synthesis of available evidence surrounding the ecology and dispersal strategies of phasmid eggs and weigh up the evidence for convergent evolution between phasmid eggs and seeds. In doing so, we highlight areas where further research is needed and discuss how the ecology of phasmid eggs may interplay with other aspects of phasmid ecology, distribution, and evolution.

Large-scale mutation in the evolution of a gene complex for cryptic coloration.

The types of mutations affecting adaptation in the wild are only beginning to be understood. In particular, whether structural changes shape adaptation by suppressing recombination or by creating new mutations is unresolved. Here, we show that multiple linked but recombining loci underlie cryptic color morphs of Timema chumash stick insects. In a related species, these loci are found in a region of suppressed recombination, forming a supergene. However, in seven species of Timema, we found that a megabase-size "supermutation" has deleted color loci in green morphs. Moreover, we found that balancing selection likely contributes more to maintaining this mutation than does introgression. Our results show how suppressed recombination and large-scale mutation can help to package gene complexes into discrete units of diversity such as morphs, ecotypes, or species.

Estimating densities of larval Salmonflies (Pteronarcys californica) through multiple pass removal of post-emergent exuvia in Colorado rivers.

Traditional methods of collecting, sorting, and identifying benthic macroinvertebrate samples are useful for stream biomonitoring and ecological studies, however, these methods are time consuming, expensive, and require taxonomic expertise. Estimating larval densities through collection of post-emergent exuvia can be a practical and time efficient alternative. We evaluated the use of multiple pass depletion techniques of the post-emergent exuvia of Pteronarcys californica to estimate larval densities at ten sites in three Colorado rivers. Exuvia density was highly correlated with both final-instar larval density (R2 = 0.90) and total larval density (R2 = 0.88) and the multiple pass removal technique performed well. Exuvia surveys found P. californica at three low density sites where benthic sampling failed to detect it. At moderate and high density sites the exuvia surveys always produced lower density estimates than benthic surveys. Multiple pass depletion estimates of exuvia proved to be an accurate and efficient technique at estimating larval densities and provided an effective alternative for traditional benthic sampling when objectives are detecting and monitoring P. californica, especially at low density sites.

The genetic mechanism of selfishness and altruism in parent-offspring coadaptation.

The social bond between parents and offspring is characterized by coadaptation and balance between altruistic and selfish tendencies. However, its underlying genetic mechanism remains poorly understood. Using transcriptomic screens in the subsocial European earwig, Forficula auricularia, we found the expression of more than 1600 genes associated with experimentally manipulated parenting. We identified two genes, Th and PebIII, each showing evidence of differential coexpression between treatments in mothers and their offspring. In vivo RNAi experiments confirmed direct and indirect genetic effects of Th and PebIII on behavior and fitness, including maternal food provisioning and reproduction, and offspring development and survival. The direction of the effects consistently indicated a reciprocally altruistic function for Th and a reciprocally selfish function for PebIII. Further metabolic pathway analyses suggested roles for Th-restricted endogenous dopaminergic reward, PebIII-mediated chemical communication and a link to insulin signaling, juvenile hormone, and vitellogenin in parent-offspring coadaptation and social evolution.

Geographical origin determines responses to salinity of Mediterranean caddisflies.

Many freshwater ecosystems worldwide, and particularly Mediterranean ones, show increasing levels of salinity. These changes in water conditions could affect abundance and distribution of inhabiting species as well as the provision of ecosystem services. In this study we conduct laboratory experiments using the macroinvertebrate Smicridea annulicornis as a model organism. Our factorial experiments were designed to evaluate the effects of geographical origin of organisms and salinity levels on survival and behavioral responses of caddisflies. The experimental organisms were captured from rivers belonging to three hydrological basins along a 450 Km latitudinal gradient in the Mediterranean region of Chile. Animals were exposed to three conductivity levels, from 180 to 1400 µS/cm, close to the historical averages of the source rivers. We measured the behavioral responses to experimental stimuli and the survival time. Our results showed that geographical origin shaped the behavioral and survival responses to salinity. In particular, survival and activity decreased more strongly with increasing salinity in organisms coming from more dilute waters. This suggests local adaptation to be determinant for salinity responses in this benthic invertebrate species. In the current scenario of fast temporal and spatial changes in water levels and salt concentration, the conservation of geographic intra-specific variation of aquatic species is crucial for lowering the risk of salinity-driven biodiversity loss.

Pressure-induced silk spinning mechanism in webspinners (Insecta: Embioptera).

The articulated appendages of arthropods are highly adaptable and potentially multifunctional, used for walking, swimming, feeding, prey capture, or other functions. Webspinners (Order Embioptera) are a paragon in this context. In contrast to other arthropods producing silk, they utilize their front feet for silk production. However, employing the same leg for alternative functions rather than for pure locomotion potentially imposes constraints and compromises. We here present morphological and experimental evidence for a "passive" pressure-induced silk spinning mechanism induced by external mechanical stimuli. Furthermore, we demonstrate that, as a consequence of the conflicting functions for their front feet, webspinners have evolved a unique style of walking that reduces the potentially problematic contact between silk ejectors and the substrate. Here we answer for the first time a long-term question within this enigmatic group of insects-how webspinners can use their front feet to spin their nanoscale silk. This knowledge may open the door for experimental studies on an artificial spinning process and for future utilization in applied fields of robotics or chemistry.

Distribution and habitat preference of Ephemeroptera and Trichoptera in subtropical mountain streams: implications for monitoring and conservation.

The assessment of biotic-habitat relationships provides key information to predict biotic responses to perturbations and important tools for river management and monitoring. This study aimed to assess the spatial distribution of Ephemeroptera and Trichoptera in mountain streams of central Argentina. We evaluated the effect of seasonality and identified the variables conditioning the abundance of the assemblages and the habitat with the highest taxonomic richness. Sampling was carried out in four streams (Carcarañá River basin) during high and low water periods. Three lotic habitats were sampled: riffles, coarse substrate runs, and fine substrate runs; and physicochemical and habitat variables were measured. We found differences in assemblage composition, among the 25 genera of Ephemeroptera and Trichoptera, at the habitat level. The most important variables affecting distribution patterns were substrate and flow type, but macroalgae and organic matter (twigs, leaves, and detritus) were also important predictors. Riffles maintained the highest richness but when considering only the Ephemeroptera taxa, fine substrate runs emerged also as an important habitat for these taxa. Our study provided valuable ecological information related to habitat preference of taxa with a key role in stream functioning and of great usefulness for the monitoring of lotic systems.

Viviparity in Two Closely Related Epizoic Dermapterans Relies on Disparate Modifications of Reproductive Systems and Embryogenesis.

Nutritional modes operating during embryonic/larval development of viviparous species range from "pure" lecitothrophy in which embryos rely solely on reserve materials (yolk spheres, lipid droplets, and glycogen particles) accumulated in the egg cytoplasm to matrotrophy in which embryos are continuously supplied with nutrients from a parental organism. Interestingly, a wide spectrum of diverse "mixed" modes employed in the embryo nourishment have also been described among viviparous species. Here, we summarize results of histochemical, ultrastructural, and biochemical analyses of reproductive systems as well as developing embryos of two closely related viviparous species of earwigs (Dermaptera), Hemimerus talpoides and Arixenia esau. These analyses clearly indicate that morphological as well as physiological modifications (adaptations) supporting viviparity and matrotrophy in Hemimerus and Arixenia, with the exception of a complex biphasic respiration, are markedly different. Most importantly, Hemimerus embryos complete their development inside terminal (largest) ovarian follicles, whereas Arixenia embryos, after initial developmental stages, are transferred to highly modified lateral oviducts, that is the uterus, where they develop until the release (birth) of larvae. The obtained results strongly suggest that viviparity in hemimerids and arixeniids had evolved independently and might therefore serve as an example of evolutionary parallelism as well as remarkable functional plasticity of insect reproduction and embryonic development.

Parallel evolution of an alpine type ecomorph in a scorpionfly: Independent adaptation to high-altitude environments in multiple mountain locations.

Elucidation of the diversification process of organisms is one of the important tasks of biology. From the viewpoint of species diversity, insects are the most successful group among the diverse organisms on earth and evolutionary adaptation is one of the important factors driving this pattern. Evolutionary adaptation is one of the important factors in the diversification of insects. One of the representative examples of environmental adaptation in insects is the shortening and loss of wings in subalpine and alpine zones. In this study, we focused on the Japanese scorpionfly, Panorpodes paradoxus. In this species, individuals that inhabit mountainous regions and subalpine zones have long wings (the "general type"), and individuals that inhabit higher altitudinal ranges have short wings (the "alpine type"). We collected samples of all Japanese Panorpodes species and one Korean Panorpodes species, and conducted molecular phylogenetic analyses of the mtDNA COI (610 bp), COII (688 bp), and 16S rRNA (888 bp) and nuDNA EF1-α (658 bp) and 28S rRNA (524 bp) regions in order to reveal the evolutionary history of the alpine type of P. paradoxus. As a result of molecular phylogenetic analyses, it was revealed that the alpine type of P. paradoxus was polyphyletic, and had evolved to become the alpine type at least twice independently at separate mountain locations. In addition, the result of divergence time estimation suggested that the alpine type is an "ecomorph", having recently adapted to low temperature habitats following mountain uplift within the Japanese Archipelago and subsequent glacial-interglacial cycles.

Attraction of the Green Lacewing Chrysoperla comanche (Neuroptera: Chrysopidae) to Yeast.

Many adult Chrysoperla comanche (Stephens) green lacewings were caught in traps baited with live yeast cultures during tests designed to catch olive fruit flies. All 13 yeast species tested were more attractive than the industry-standard dried torula yeast (Cyberlindnera jadinii; syn. Candida utilis). Live C. jadinii culture attracted significantly more lacewings than the inactive dried-pellet form of the same yeast species, demonstrating that volatiles from live yeast cultures attract adults of this lacewing. Odor profiles for two of the highly active yeasts tested herein (Lachancea thermotolerans and Solicoccozyma terrea) were similar to that for Metschnikowia pulcherrima, a yeast species isolated earlier from the gut diverticulum of Chrysoperla rufilabris. A new Metschnikowia species (M. chrysoperlae), along with two new Candida spp. that were recently realigned to one of the Metschnikowia clades (M. picachoensis and M. pimensis), were also identified from the diverticulum of C. comanche. Thus, one clade of Metschnikowia yeasts that commonly occur in floral nectar appears to exhibit mutualistic symbioses with Chrysoperla green lacewings. Both male and female C. comanche adults were attracted in the present study, and we speculate that males have exploited this symbiosis by offering Metschnikowia-laden regurgitant, including attractive volatiles, to females ('mating trophallaxis') as a nuptial gift.

Pest and natural enemy: how the fat bodies of both the southern armyworm Spodoptera eridania and the predator Ceraeochrysa claveri react to azadirachtin exposure.

The effects of biopesticides on insects can be demonstrated by morphological and ultrastructural tools in ecotoxicological analysis. Azadirachtin-based products are widely used as biopesticides, affecting numerous insect populations. Through morphological biomarkers, this study aimed to characterize the fat bodies of both the southern armyworm Spodoptera eridania and the predator Ceraeochrysa claveri after chronic exposure to azadirachtin. Larvae of S. eridania and C. claveri were fed with fresh purple lettuce leaves (Lactuca sativa) and egg clusters of Diatraea saccharalis treated with azadirachtin solution of 6 mg active ingredient (a.i.)/L and 18 mg a.i./L for 7 days, respectively. The biological data showed a significant reduction in survival and body mass in S. eridania and cytotoxic effects in the parietal and perivisceral fat bodies in both species. Ultrastructural cell damage was observed in the trophocytes of both species such as dilated cisternae of the rough endoplasmic reticulum and swollen mitochondria. Trophocytes of S. eridania and C. claveri of the parietal and perivisceral layers responded to those injuries by different cytoprotective and detoxification means such as an increase in the amount of cytoplasmic granules containing calcium, expression of heat shock protein (HSP)70/HSP90, and development of the smooth endoplasmic reticulum. Despite all the different means of cytoprotection and detoxification, they were not sufficient to recover from all the cellular damages. Azadirachtin exhibited an excellent performance for the control of S. eridania and a moderate selectivity for the predator C. claveri, which presents better biological and cytoprotective responses to chronic exposure to azadirachtin.

Evolutionary history of Polyneoptera and its implications for our understanding of early winged insects.

Polyneoptera represents one of the major lineages of winged insects, comprising around 40,000 extant species in 10 traditional orders, including grasshoppers, roaches, and stoneflies. Many important aspects of polyneopteran evolution, such as their phylogenetic relationships, changes in their external appearance, their habitat preferences, and social behavior, are unresolved and are a major enigma in entomology. These ambiguities also have direct consequences for our understanding of the evolution of winged insects in general; for example, with respect to the ancestral habitats of adults and juveniles. We addressed these issues with a large-scale phylogenomic analysis and used the reconstructed phylogenetic relationships to trace the evolution of 112 characters associated with the external appearance and the lifestyle of winged insects. Our inferences suggest that the last common ancestors of Polyneoptera and of the winged insects were terrestrial throughout their lives, implying that wings did not evolve in an aquatic environment. The appearance of the first polyneopteran insect was mainly characterized by ancestral traits such as long segmented abdominal appendages and biting mouthparts held below the head capsule. This ancestor lived in association with the ground, which led to various specializations including hardened forewings and unique tarsal attachment structures. However, within Polyneoptera, several groups switched separately to a life on plants. In contrast to a previous hypothesis, we found that social behavior was not part of the polyneopteran ground plan. In other traits, such as the biting mouthparts, Polyneoptera shows a high degree of evolutionary conservatism unique among the major lineages of winged insects.

Potential Effects of Future Climate Changes on Brazilian Cool-Adapted Stoneflies (Insecta: Plecoptera).

The continuous pursuit of welfare and economic development through the exploitation of natural resources by human societies consequently resulted in the ongoing process of climate change. Changes in the distribution of species towards the planet's poles and mountain tops are some of the expected to biological consequences of this process. Here, we assessed the potential effects of future climate change on four cool-adapted Gripopterygidae (Insecta: Plecoptera) species [Gripopteryx garbei Navás 1936, G. cancellata (Pictet 1841), Tupiperla gracilis (Burmeister 1839), and T. tessellata (Brauer 1866)] from Southeastern Brazilian Atlantic forest. As species adapted to cold conditions, in the future scenarios of climate change, we expected these organisms to shrink/change their distributions ranges towards areas with suitable climatic conditions in Southern Brazilian regions, when compared with their predicted distributions in present climatic conditions. We used seven principal components derived from 19 environmental variables from Worldclim database for the present scenario and also seven principal components obtained from 17 different Atmosphere-Ocean Global Circulation Models (AOGCMs), considering the most severe emission scenario for green-house gases to predict the species' distributions. Depending on the climatic scenario considered, there were polewards distribution range changes of the species. Additionally, we also observed an important decrease in the amount of protected modeled range for the species in the future scenarios. Considering that this Brazilian region may become hotter in the future and have its precipitation regime changed, as observed in the severe 2013-2014 drought, we believe these species adapted to high altitudes will be severely threatened in the future.