Insect Size Research Articles

A comparative study of body size evolution in moths: evidence of correlated evolution with feeding and phenology-related traits.

Interspecific variation in body size is one of the most popular topics in comparative studies. Despite recent advances, little is known about the patterns and processes behind the evolution of body size in insects. Here, we used a robust data set comprising all geometrid moth species occurring in Northern Europe to examine the evolutionary associations involving body size and several life-history traits under an explicitly phylogenetic framework. We provided new insights into the interactive effects of life-history traits on body size and evidence of correlated evolution. We further established the sequence of trait evolution linking body size with the life-history traits correlated with it. We found that most (but not all) of the studied life-history traits, to some extent, influenced interspecific variation in body size, but interactive effects were uncommon. Both bi- and multivariate phylogenetic analyses indicated that larger species tend to be nocturnal flyers, overwinter in the larval stage, feed on the foliage of trees rather than herbs, and have a generalist feeding behaviour. We found evidence of correlated evolution involving body size with overwintering stage, host-plant growth form, and dietary specialization. The examination of evolutionary transitions within the correlated evolution models signalled that overwintering as larvae commonly preceded the evolution of large sizes, as did feeding on tree foliage and the generalist feeding behaviour. By showing that both body size and all life-history traits correlated with it evolve at very slow rates, we caution against uncritical attempts to propose causal explanations for respective associations based on contemporary ecological settings.

Effect of temperature on sexual size dimorphism during the developmental period in the broad-horned flour beetle

Adult size in numerous insects is strongly dependent on temperature. In several cases, a temperature–size rule is observed in which developmental temperature and adult size tradeoff. Although several previous studies have demonstrated the temperature–size rule, only a few have explored the relationship between developmental temperature and weapon traits or sexual size dimorphism. This study was conducted to investigate the size of the broad-horned flour beetle Gnatocerus cornutus when it was developed under different temperatures. G. cornutus males possess weapon traits for male–male combat and exhibit sexual size dimorphism in other morphological traits. Results showed that male weapon size and body size complied with the temperature–size rule. Furthermore, the extent of sex dimorphism in genae width, a weapon-supportive trait, were larger at lower temperatures. Our findings suggest that the temperature–size rule also influences the size of sexual traits.

Reduced size in a montane butterfly at its warm range boundaries

Abstract Variation in insect size is often related to temperature during development and may affect the persistence of populations under future climate warming if smaller individuals have reduced fitness. Montane species are particularly vulnerable to climate‐driven local extinctions due to range retractions at their warm range margins, and so we examined spatial and temporal variation in body size in the butterfly Erebia epiphron (Lepidoptera: Nymphalidae) in the United Kingdom, where it is restricted to two montane regions in England and Scotland. We examined spatial and temporal variation in body size in relation to temperature. We sampled 19 populations (6–15 individuals per population) in England and Scotland between 2018 and 2019 spanning elevations from 380 to 720 m and examined museum specimens collected between 1890 and 1980. We examined individual body size (forewing length) and its relationship with the local temperature of sites, as well as temporal variation in body size over the last century in relation to the temperature during larval development. The forewing lengths of field‐collected individuals in England were on average 7%–8% smaller than in Scotland (England, mean = 14.9 mm, Scotland, mean = 15.9 mm), and warmer sites also had smaller individuals (0.13 mm reduction in wing length per 1°C increase in local site mean temperature). However, we found no effect of temporal temperature variation on body size changes during larval development. The observed smaller body size in English populations could have impacts on fecundity and dispersal ability. Future work should seek to understand the life‐cycle lengths, genetics and phenotypic plasticity of these two populations to evaluate potential explanations for regional differences.

A strategy for improving silk yield and organ size in silk-producing insects.

Insect silks possess excellent biodegradability, biocompatibility and mechanical properties, and have numerous applications in biomedicine and tissue engineering. However, the application of silk fiber is hindered by its limited supply, especially from non-domesticated insects. In the present study, the silk yield and organ size of Bombyx mori were significantly improved through genetic manipulation of the target of rapamycin complex 1 (TORC1) pathway components. Silk protein synthesis and silk gland size were decreased following rapamycin treatment, inhibiting the TORC1 signaling pathway both in vivo and ex vivo. The overexpression of posterior silk gland-specific Rheb and BmSLC7A5 improved silk protein synthesis and silk gland size by activating the TORC1 signaling pathway. Silk yield in BmSLC7A5-overexpression silkworms was significantly increased by approximately 25%. We have demonstrated that the TORC1 signaling pathway is involved in the transcription and translation of silk genes and transcriptional activators via phosphorylation of p70 S6 kinase 1 and 4E-binding protein 1. Our findings present a strategy for increasing silk yield and organ size in silk-producing insects.

Monitoring insect numbers and biodiversity with a vertical-beam entomological radar.

Concerns about perceived widespread declines in insect numbers have led to recognition of a requirement for long-term monitoring of insect biodiversity. Here we examine whether an existing, radar-based, insect monitoring system developed for research on insect migration could be adapted to this role. The radar detects individual larger (greater than 10 mg) insects flying at heights of 150-2550 m and estimates their size and mass. It operates automatically and almost continuously through both day and night. Accumulation of data over a 'half-month' (approx. 15 days) averages out weather effects and broadens the source area of the wind-borne observation sample. Insect counts are scaled or interpolated to compensate for missed observations; adjustment for variation of detectability with range and insect size is also possible. Size distributions for individual days and nights exhibit distinct peaks, representing different insect types, and Simpson and Shannon-Wiener indices of biodiversity are calculated from these. Half-month count, biomass and index statistics exhibit variations associated with the annual cycle and year to year changes that can be attributed to drought and periods of high rainfall. While species-based biodiversity measures cannot be provided, the radar's capacity to estimate insect biomass over a wide area indicates utility for tracking insect population sizes. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Enhancing Insect Sound Classification Using Dual-Tower Network: A Fusion of Temporal and Spectral Feature Perception

In the modern field of biological pest control, especially in the realm of insect population monitoring, deep learning methods have made further advancements. However, due to the small size and elusive nature of insects, visual detection is often impractical. In this context, the recognition of insect sound features becomes crucial. In our study, we introduce a classification module called the “dual-frequency and spectral fusion module (DFSM)”, which enhances the performance of transfer learning models in audio classification tasks. Our approach combines the efficiency of EfficientNet with the hierarchical design of the Dual Towers, drawing inspiration from the way the insect neural system processes sound signals. This enables our model to effectively capture spectral features in insect sounds and form multiscale perceptions through inter-tower skip connections. Through detailed qualitative and quantitative evaluations, as well as comparisons with leading traditional insect sound recognition methods, we demonstrate the advantages of our approach in the field of insect sound classification. Our method achieves an accuracy of 80.26% on InsectSet32, surpassing existing state-of-the-art models by 3 percentage points. Additionally, we conducted generalization experiments using three classic audio datasets. The results indicate that DFSM exhibits strong robustness and wide applicability, with minimal performance variations even when handling different input features.

A pipeline contributes to efficient identification of salivary proteins in short-headed planthopper, Epeurysa nawaii

Saliva, an oral secretion primarily originating from salivary glands (SGs), exert critical roles in the ongoing evolutionary interaction between insects and plants. However, identifying insect salivary components poses challenges due to the tiny size of insects, low secretion amounts, and the propensity for degradation after secretion. In this study, we developed a transcriptome-based approach to comprehensively analyze the salivary proteins of the short-headed planthopper, Epeurysa nawaii, a species with unique feeding habits on bamboo. A total of 165 salivary proteins were identified, with 114 secretory genes highly and specifically expressed in SGs. Consistent with most phloem-feeding insects, digestive enzymes, calcium-binding proteins, oxidoreductases, and a few previously reported salivary effectors were ubiquitously distributed in E. nawaii saliva. However, we also identified a substantial portion of salivary proteins exhibiting taxonomy specificity, including 60 E. nawaii-specific and 62 Delphacidae-specific proteins. These taxonomy-restricted proteins potentially play a role in insect adaptation to specific host plants. Our study provides an efficient pipeline for salivary protein identification and serves as a valuable resource for the functional characterization of effectors.

Improving accuracy of age estimates for insect evidence-calibration of physiological age at emergence (k) using insect size but without "k versus size" model.

Postmortem interval may be estimated based on the age of insect evidence collected on a death scene. Reference data that are used in such estimation frequently comprise thermal summation constant (i.e. k), which is equal to the insect age upon completion of immature development expressed in accumulated degree-days or degree-hours (ADD or ADH). Essentially, k is a central point of an insect group and it may poorly represent insect evidence that is near the limits of variation for the group. Accordingly, it was postulated to calibrate k for particular insect evidence and insect size and sex were found to be useful for this purpose in some of the species. However, the calibration is only possible by using the model that correlates k with insect size. Since very few such models were published, this lack of data limits the calibration of k in forensic casework. In this article, we develop a formula that is useful for the calibration of k without the use of "k versus size" model (and related datasets). The formula uses k from the general thermal summation model for a species (with its standard error), size range for the species (retrieved from entomology literature), and size measurements for particular insect evidence. The calibration of k with the formula was validated using the Creophilus maxillosus (Coleoptera: Staphylinidae) and Necrodes littoralis (Coleoptera: Silphidae) datasets. It was particularly useful while analyzing unusually small and large insects, in case of which the formula reduced the inaccuracy of k from the general model on average by ~25 ADD in C. maxillosus and ~40 ADD in N. littoralis. We discuss the limitations and prospects of the calibration protocol that employs the formula.

STARdbi: A pipeline and database for insect monitoring based on automated image analysis

Insects are highly abundant and diverse, and play major roles in ecosystem functions. Monitoring of insect populations is key to their sustainable management. However, the labor and expertise needed to identify insects, and the challenges of archiving the wealth of data collected in monitoring programs, often limit these efforts. We describe a pipeline to reduce the barriers associated with curating and mining big data of insect biodiversity. The pipeline, STARdbi, includes capturing flying insects with sticky traps, scanning the traps, storing the trap-images in a public database with a web-based interface, and applying machine learning models to extract information from the images. To illustrate the insights that can be gained from STARdbi, we describe two case studies. One of them involves monitoring of circadian activity patterns of grain pests and of their natural enemies, and the other compares insect abundance, biomass and size distributions between agricultural and semi-natural habitats. We invite the community of insect ecologists to contribute to the STARdbi database, and to use its image analysis tools to address diverse ecological and evolutionary questions.

Spectral composition of light-emitting diodes impacts aquatic and terrestrial invertebrate communities with potential implications for cross-ecosystem subsidies.

Resource exchanges in the form of invertebrate fluxes are a key component of aquatic-terrestrial habitat coupling, but this interface is susceptible to human activities, including the imposition of artificial light at night. To better understand the effects of spectral composition of light-emitting diodes (LEDs)-a technology that is rapidly supplanting other lighting types-on emergent aquatic insects and terrestrial insects, we experimentally added LED fixtures that emit different light spectra to the littoral zone and adjacent riparian habitat of a pond. We installed four replicate LED treatments of different wavelengths (410, 530 and 630 nm), neutral white (4000 k) and a dark control, and sampled invertebrates in both terrestrial and over-water littoral traps. Invertebrate communities differed among light treatments and between habitats, as did total insect biomass and mean individual insect size. Proportional allochthonous biomass was greater in the riparian habitat and among some light treatments, demonstrating an asymmetrical effect of differently coloured LEDs on aquatic-terrestrial resource exchanges. Overall, our findings demonstrate that variation in wavelength from LEDs may impact the flux of resources between systems, as well as the communities of insects that are attracted to particular spectra of LED lighting, with probable implications for consumers. This article is part of the theme issue 'Light pollution in complex ecological systems'.

The response to fire by two eusocial bee species

Abstract Fire is a common worldwide disturbance that affected the evolution of life and shaped plant and animal diversity; this is especially applicable in the Mediterranean. In this study, we explore the response to fire of two eusocial bee species of Halictidae, namely, Lasioglossum marginatum (Brullé, 1832) and Lasioglossum malachurum (Kirby, 1802), as to their population number, body size and pollination specialisation. The study was carried out on Chios Island, Greece, during the first 3 years after a wildfire (2013–2015). Fire effect was examined from a spatial viewpoint by comparing unburnt with burnt sites and from a temporal one by comparing populations among different postfire years. We found that L. malachurum populations increased with time after fire, while they were more abundant in the burnt sites during the second postfire year. In contrast, L. marginatum populations decreased in both burnt and unburnt sites, which suggests that the decline was not an effect of the fire alone. Insect body size differed among years for both species, with L. marginatum individuals being smaller in the burnt sites compared to the unburnt, albeit only in the first postfire year. Finally, both species show high partner fidelity but to different plant partners. Our results underscore the importance of studies focusing on the effects of a disturbance at the intraspecific level. However long‐term monitoring is required, especially in the context of the global pollinator crisis and the more frequent and severe wildfires as an effect of climate change.

A method for estimating colony size using queen fecundity in termites under field conditions.

Colony size in social insects is one of the most important factors in shaping their self-organized system. It affects a wide variety of traits such as foraging and defense strategies, social immune responses, the degree of polymorphism, and reproductive output. However, colony size estimation of subterranean termites in the field has been challenging, due to their extremely cryptic biology and multiple site-nesting behavior. Since natural selection favors workers that maximize the number of their siblings, the amount of egg production may reflect the number of workers in the colony. Here, we report a method for inferring colony size in the field using total egg production in each colony from a subterranean termite, Reticulitermes speratus. Our investigation of field colonies revealed that the body weight of queens reaches a peak and had the largest variance in June and July and accurately predicts the number of eggs laid by the queen per 24 h. Using laboratory-reared colonies, we found that the total egg production in each colony is proportional to the number of workers. We also estimated the colony size of 198 field colonies and found that the median and maximum colony size was 24,500 and 451,800 workers per colony. The method for inferring colony size presented here may also be applicable to termite species with a clear seasonality in egg production. The colony size estimate will contribute to understanding the life history strategies and social systems of termites.

Aquatic insect accumulation of uranium at spring outflows in the Grand Canyon region as influenced by aqueous and sediment geochemistry and biological factors: implications for monitoring.

Potential adverse ecological effects of expanded uranium (U) mining within the Grand Canyon region motivated studies to better understand U exposure and risk to endemic species. This study documents U exposures and analyzes geochemical and biological factors affecting U bioaccumulation at spring-fed systems within the Grand Canyon region. The principal objective was to determine if aqueous U was broadly indicative of U accumulated by insect larvae, a dominate fauna. Analyses focused on three widely distributed taxa: Argia sp. (a predatory damselfly), Culicidae (suspension feeding mosquitos), and Limnephilus sp. (a detritivorous caddisfly). The study showed that U accumulated by aquatic insects (and periphyton) generally correlated positively with total dissolved U, although correlations were strongest when based on modeled concentrations of the U-dicarbonato complex, UO2(CO3)2-2, and UO2(OH)2. Sediment metal concentration was a redundant indicator of U bioaccumulation. Neither insect size or U in the gut content of Limnephilus sp. substantially affected correlations between aqueous U and whole-body U concentrations. However, in Limnephilus sp., the gut and its content contained large quantities of U. Estimates of the sediment burden in the gut indicated that sediment was a minor source of U mass but contributed substantially to the total insect weight. As a result, whole-body U concentration would tend to vary inversely with the sediment burden of the gut. The correlations between aqueous U and bioaccumulated U provide an initial relational baseline against which newly acquired data could be evaluated for changes in U exposure during and after mining operations.

Early-life food stress hits females harder than males in insects: A meta-analysis of sex differences in environmental sensitivity.

Fitness consequences of early-life environmental conditions are often sex-specific, but corresponding evidence for invertebrates remains inconclusive. Here, we use meta-analysis to evaluate sex-specific sensitivity to larval nutritional conditions in insects. Using literature-derived data for 85 species with broad phylogenetic and ecological coverage, we show that females are generally more sensitive to food stress than males. Stressful nutritional conditions during larval development typically lead to female-biased mortality and thus increasingly male-biased sex ratios of emerging adults. We further demonstrate that the general trend of higher sensitivity to food stress in females can primarily be attributed to their typically larger body size in insects and hence higher energy needs during development. By contrast, there is no consistent evidence of sex-biased sensitivity in sexually size-monomorphic species. Drawing conclusions regarding sex-biased sensitivity in species with male-biased size dimorphism remains to wait for the accumulation of relevant data. Our results suggest that environmental conditions leading to elevated juvenile mortality may potentially affect the performance of insect populations further by reducing the proportion of females among individuals reaching reproductive age. Accounting for sex-biased mortality is therefore essential to understanding the dynamics and demography of insect populations, not least importantly in the context of ongoing insect declines.

Microbial diversity in stingless bee gut is linked to host wing size and influenced by the environment

Stingless bees are important social corbiculate bees, fulfilling critical pollination roles in many ecosystems. However, their gut microbiota, particularly the fungal communities associated with them, remains inadequately characterised. This knowledge gap hinders our understanding of bee gut microbiomes and their impacts on the host fitness. We collected 121 samples from two species, Tetragonula carbonaria and Austroplebeia australis across 1200 km of eastern Australia. We characterised their gut microbiomes and investigated potential correlations between bee gut microbiomes and various geographical and morphological factors. We found their core microbiomes consisted of the abundant bacterial taxa Snodgrassella, Lactobacillus and Acetobacteraceae, and the fungal taxa Didymellaceae, Monocilium mucidum and Aureobasidium pullulans, but variances of their abundances among samples were large. Furthermore, gut bacterial richness of T. carbonaria was positively correlated to host forewing length, an established correlate to body size and fitness indicator in insects relating to flight capacity. This result indicates that larger body size/longer foraging distance of bees could associate with greater microbial diversity in gut. Additionally, both host species identity and management approach significantly influenced gut microbial diversity and composition, and similarity between colonies for both species decreased as the geographic distance between them increased. We also quantified the total bacterial and fungal abundance of the samples using qPCR analyses and found that bacterial abundance was higher in T. carbonaria compared to A. australis, and fungi were either lowly abundant or below the threshold of detection for both species. Overall, our study provides novel understanding of stingless bee gut microbiomes over a large geographic span and reveals that gut fungal communities likely not play an important role in host functions due to their low abundances.

Lyric Machines: Insects in Seventeenth-Century Poetry

Abstract This essay addresses the intriguing frequency of insect lyrics in seventeenth-century English poetry. While dramatic developments in the scientific and artistic regimes, including the invention of the microscope and the rise of still-life painting, undoubtedly played a role in this proliferation of entomological texts, this essay suggests that the figure of the insect was also deployed by early modern poets to probe the formal and imaginative potentialities of lyric, a genre that assumed a prominence over the course of the seventeenth century. The diminutive size, brief lifespan, and intricate anatomy of insects resonate with the formal qualities of lyric as short, compressed, and technically demanding objects of poetic matter, and the ambivalent attitudes to insects embody the contradictions surrounding the early modern idea of lyric. By using the life of fleas, glowworms, flies, bees, and grasshoppers to think through questions of lyric ontology, early modern poets from Donne to Killigrew redraw the familiar contours of the lyric genre as a contested territory of fascination and disgust with insect poiesis caught between human and animal, nature and machine, and life and death.

Bumblebee cognitive abilities are robust to changes in colony size

Eusocial insect colonies act as a superorganism, which can improve their ability to buffer the negative impact of some anthropogenic stressors. However, this buffering effect can be affected by anthropogenic factors that reduce their colony size. A reduction in colony size is known to negatively affect several parameters like brood maintenance or thermoregulation, but the effects on behaviour and cognition have been largely overlooked. It remains unclear how a sudden change in group size, such as that which might be caused by anthropogenic stressors, affects individual behaviour within a colony. In this study, the bumblebee Bombus terrestris was used to study the effect of social group size on behaviour by comparing the associative learning capabilities of individuals from colonies that were unmanipulated, reduced to a normal size (a colony of 100 workers) or reduced to a critically low but functional size (a colony of 20 workers). The results demonstrated that workers from the different treatments performed equally well in associative learning tasks, which also included no significant differences in the learning capacity of workers that had fully developed after the colony size manipulation. Furthermore, we found that the size of workers had no impact on associative learning ability. The learning abilities of bumblebee workers were thus resilient to the colony reduction they encountered. Our study is a first step towards understanding how eusocial insect cognition can be impacted by drastic reductions in colony size.Significance statementWhile anthropogenic stressors can reduce the colony size of eusocial insects, the impact of this reduction is poorly studied, particularly among bumblebees. We hypothesised that colony size reduction would affect the cognitive capacity of worker bumblebees as a result of fewer social interactions or potential undernourishment. Using differential conditioning, we showed that drastic reductions in colony size have no effect on the associative learning capabilities of the bumblebee Bombus terrestris and that this was the same for individuals that were tested just after the colony reduction and individuals that fully developed under the colony size reduction. We also showed that body size did not affect learning capabilities. This resilience could be an efficient buffer against the ongoing impacts of global change.

3D-Printed High-Frequency Dielectric Elastomer Actuator toward Insect-Scale Ultrafast Soft Robot

Insects with small and light bodies possess the capability of agile and fast movement in a small space. Inspired by nature, an insect-like soft robot may update the strategies in many scenarios like exploration, rescue, etc. However, the design and mass manufacture of soft robots combining insect size, fast mobility, good robustness, and impact-perception capability still present great engineering challenges. Herein, we report an insect-scale (15 mm body length (BL), 450 mg body weight) and ultrafast (∼4.0 BL s–1) soft robot. The remarkable motion performance is attributed to the high-frequency (760 Hz) operation as well as the long lifetime (>one million cycles) of its artificial muscle, which is a coil dielectric elastomer actuator (DEA) made by multimaterial coaxial three-dimensional printing with well-designed highly elastic materials and 5-inlets nozzle structure. The current robot is not only the smallest and fastest among the reported DEA-driven robots but also obtains high robustness, good environmental adaptability, and impact-perception capability: It can run on various grounds and complex paths, climb inside small pipes, work in robot swarms, and sustain and perceive the impact of the external environment.

Sex- and size-dependent variation in wing morphology of the cuckoo wasp Trichrysis cyanea (Hymenoptera: Chrysididae)

Intraspecific variation and sexual dimorphism in wing shape and size is common in winged insects. The exact patterns, however, differ among taxa and are related to the selection pressure acting on specific traits. Cuckoo wasps (Chrysdidae) are hymenopterans that have evolved a complex parasitoid or cleptoparasitic life-style. For the first time, we studied the intraspecific variation in wing shape and size in the model species, Trichysis cyanea, a common Palearctic cuckoo wasp. This involved geometric morphometrics combined with a novel, non-invasive way of obtaining images, to study the shape and size of the forewings of males and females. We found clear sexual dimorphism in both wing shape and size, possibly related to fecundity (as wing size typically correlates with body size in many insects) and increased manoeuvrability of females for searching for their host's nesting holes. This study increases our knowledge of the biology of T. cyanea in terms of a better understanding of the evolution of parasitism and corresponding adaptations in this hymenopteran family.

RNA interference knockdown of insulin receptor inhibits ovarian development in Chilo suppressalis.

The nutritional signaling pathway regulates an insect's size, development, and lifespan, as well as playing a vital role in reproduction. The insulin/insulin-like growth factor signaling (IIS) pathway plays a key role in the nutrition signaling pathway. As an integral component of the IIS pathway, insulin receptor (InR), a receptor tyrosine kinase, plays a role in the insulin pathway by controlling reproduction in many insect species. However, the precise molecular function of InR in non-model insect reproduction is poorly understood. In our study, Chilo suppressalis, a well-known rice pest, was used as a molecular system to determine the role of InR in insect reproduction. Sequencing the InR gene of C. suppressalis, comparing the amino acid sequence-specific structure, and constructing a phylogenetic tree revealed that this gene has four main domains: ligand binding L domain, Furin-like region, fibronectin type III domains, and Tyrosine kinase catalytic domain, which were all highly conserved in insects. By characterizing the spatiotemporal expression profile of InR in different developmental stages and tissues, we found that InR gene expression was highest on the 3-day old in female pupae, 6th instar larvae, and fat body on the 6-day old in female pupae. InR gene expression may promote the molting and pupation of larvae and play a role in reproduction in the fat body. Furthermore, the RNA interference knockdown of InR dramatically reduced yolk deposition and blocked oocyte maturation. After suppression of InR, the expression of several other genes fluctuated to varying degrees. In conclusion, InR is vital to reproduction and is expected to become a new target for pest management.