Silicon applications in rice plants alter the stylet probing behaviors of Glyphepomis spinosa (Hemiptera: Pentatomidae).

The stink bug Glyphepomis spinosa Campos & Grazia (Hemiptera: Pentatomidae) is a potential rice pest in Brazil. This study evaluates the interaction between silicon sources and 3 rice cultivars (BRS Esmeralda, Canela de Ferro, and IRGA 417) and examines how increasing silicon levels affect the stylet probing behavior of G. spinosa. The experiment was set up in a completely randomized design with a 3 × 3 factorial scheme (silicon sources: calcium silicate, potassium silicate, a control, and 3 rice cultivars). Fertilizing rice plants with Si altered the probing behavior of the stink bug G. spinosa. The cultivar interaction by Si source was significant in a few variables. This was evidenced by longer periods without ingestion, prolonged time to the first stylet probe (initial probing), and less time spent in cellular maceration. This result supports the use of electropenetrography as a tool to evaluate resistance inducers in plants.

A synopsis of the genus Cylindrostethus Fieber 1861 (Insecta: Hemiptera: Gerridae).

Cylindrostethus Fieber, 1861 is one of the most striking genera of water striders (Insecta: Hemiptera: Gerridae) and has Pantropical distribution. Members of this group can be recognized by the very long, cylindrical body; the short antennomere IV; the short labium not reaching the mesosternum; and by characteristics of the abdomen of males and females. Although Neotropical representatives of the genus have been revised, there are pending taxonomic issues related to this fauna, and that of the Eastern Hemisphere has been barely studied in recent years. Here, we present a short note about the authorship of Cylindrostethus, an updated key to all species of the genus, a new synonymy, and the description of a previously unknown macropterous male of C. hungerfordi Drake and Harris.

Elucidating the ecophysiology of soybean pod-sucking stinkbug Riptortus pedestris (Hemiptera: Alydidae) based on de novo genome assembly and transcriptome analysis.

Food security is important for the ever-growing global population. Soybean, Glycine max (L.) Merr., is cultivated worldwide providing a key source of food, protein and oil. Hence, it is imperative to maintain or to increase its yield under different conditions including challenges caused by abiotic and biotic stresses. In recent years, the soybean pod-sucking stinkbug Riptortus pedestris has emerged as an important agricultural insect pest in East, South and Southeast Asia. Here, we present a genomics resource for R. pedestris including its genome assembly, messenger RNA (mRNA) and microRNA (miRNA) transcriptomes at different developmental stages and from different organs. As insect hormone biosynthesis genes (genes involved in metamorphosis) and their regulators such as miRNAs are potential targets for pest control, we analyzed the sesquiterpenoid (juvenile) and ecdysteroid (molting) hormone biosynthesis pathway genes including their miRNAs and relevant neuropeptides. Temporal gene expression changes of these insect hormone biosynthesis pathways were observed at different developmental stages. Similarly, a diet-specific response in gene expression was also observed in both head and salivary glands. Furthermore, we observed that microRNAs (bantam, miR-14, miR-316, and miR-263) of R. pedestris fed with different types of soybeans were differentially expressed in the salivary glands indicating a diet-specific response. Interestingly, the opposite arms of miR-281 (-5p and -3p), a miRNA involved in regulating development, were predicted to target Hmgs genes of R. pedestris and soybean, respectively. These observations among others highlight stinkbug's responses as a function of its interaction with soybean. In brief, the results of this study not only present salient findings that could be of potential use in pest management and mitigation but also provide an invaluable resource for R. pedestris as an insect model to facilitate studies on plant-pest interactions.

Binding properties of chemosensory protein 4 in Riptortus pedestris to aggregation pheromones.

In insects, chemosensory proteins (CSPs) play an important role in the perception of the external environment and have been widely used for protein-binding characterization. Riptortus pedestris has received increased attention as a potential cause of soybean staygreen syndrome in recent years. In this study, we found that RpedCSP4 expression in the antennae of adult R. pedestris increased with age, with no significant difference in expression level observed between males and females, as determined through quantitative real-time polymerase chain reaction (qRT-PCR). Subsequently, we investigated the ability of RpedCSP4 to bind various ligands (five aggregated pheromone components and 13 soybean volatiles) using a prokaryotic expression system and fluorescence competitive binding assays. We found that RpedCSP4 binds to three aggregated pheromone components of R. pedestris, namely, ((E)-2-hexenyl (Z)-3-hexenoate (E2Z3), (E)-2-hexenyl (E)-2-hexenoate (E2E2), and (E)-2-hexenyl hexenoate (E2HH)), and that its binding capacities are most stable under acidic condition. Finally, the structure and protein-ligand interactions of RpedCSP4 were further analyzed via homology modeling, molecular docking, and targeted mutagenesis experiments. The L29A mutant exhibited a loss of binding ability to these three aggregated pheromone components. Our results show that the olfactory function of RpedCSP4 provides new insights into the binding mechanism of RpedCSPs to aggregation pheromones and contributes to discover new target candidates that will provide a theoretical basis for future population control of R. pedestris.

Begomovirus Transmission to Tomato Plants Is Not Hampered by Plant Defenses Induced by Dicyphus hesperus Knight.

Plants can respond to insect infestation and virus infection by inducing plant defenses, generally mediated by phytohormones. Moreover, plant defenses alter host quality for insect vectors with consequences for the spread of viruses. In agricultural settings, other organisms commonly interact with plants, thereby inducing plant defenses that could affect plant-virus-vector interactions. For example, plant defenses induced by omnivorous insects can modulate insect behavior. This study focused on tomato yellow leaf curl virus (TYLCV), a plant virus of the family Geminiviridae and genus Begomovirus. It is transmitted in a persistent circulative manner by the whitefly Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae), posing a global threat to tomato production. Mirids (Hemiptera: Miridae) are effective biological control agents of B. tabaci, but there is a possibility that their omnivorous nature could also interfere with the process of virus transmission. To test this hypothesis, this study first addressed to what extent the mirid bug Dicyphus hesperus Knight induces plant defenses in tomato. Subsequently, the impact of this plant-omnivore interaction on the transmission of TYLCV was evaluated. Controlled cage experiments were performed in a greenhouse setting to evaluate the impact of mirids on virus transmission and vector acquisition by B. tabaci. While we observed a reduced number of whiteflies settling on plants exposed to D. hesperus, the plant defenses induced by the mirid bug did not affect TYLCV transmission and accumulation. Additionally, whiteflies were able to acquire comparable amounts of TYLCV on mirid-exposed plants and control plants. Overall, the induction of plant defenses by D. hesperus did not influence TYLCV transmission by whiteflies on tomato.

Chromosome-level genome assembly of the predatory stink bug Arma custos.

The stink bug Arma custos (Hemiptera: Pentatomidae) is a predatory enemy successfully used for biocontrol of lepidopteran and coleopteran pests in notorious invasive species. In this study, a high-quality chromosome-scale genome assembly of A. custos was achieved through a combination of Illumina sequencing, PacBio HiFi sequencing, and Hi-C scaffolding techniques. The final assembled genome was 969.02 Mb in size, with 935.94 Mb anchored to seven chromosomes, and a scaffold N50 length of 135.75 Mb. This genome comprised 52.78% repetitive elements. The detected complete BUSCO score was 99.34%, indicating its completeness. A total of 13,708 protein-coding genes were predicted in the genome, and 13219 of them were annotated. This genome provides an invaluable resource for further research on various aspects of predatory bugs, such as biology, genetics, and functional genomics.

Identification and expression patterns of somatic piRNAs and PIWI genes in Riptortus pedestris (Hemiptera: Alydidae).

RNA interference (RNAi)-based gene silencing is a feasible and sustainable technology for the management of hemipteran pests by double-stranded RNA involvement, including small-interfering RNA, microRNA, and Piwi-interacting RNA (piRNA) pathways, that may help to decrease the usage of chemical insecticides. However, only a few data are available on the somatic piRNAs and their biogenesis genes in Riptortus pedestris, which serves as a significant pest of soybean (Glycine max). In this study, two family members of the PIWI gene were identified and characterized in R. pedestris, containing Argonaute3 (RpAgo3) and Aubergine (RpAub) genes with conserved protein domains, and their clusters were validated by phylogenetic analysis. In addition, they were widely expressed in all developmental stages of the whole body of R. pedestris and had lower expression levels in R. pedestris guts under different rearing conditions based on previous transcriptome sequencing. Furthermore, abundant clean reads were filtered to a total number of 45,998 piRNAs with uridine bias at the first nucleotide (nt) position and 26-32 nt in length by mapping onto the reference genome of R. pedestris according to our previous whole-transcriptome sequencing. Finally, our data revealed that gut bacterial changes were significantly positively or negatively associated with differentially expressed piRNAs among the five comparison groups with Pearson correlation analysis. In conclusion, these findings paved new avenues for the application of RNAi-based biopesticides for broad-spectrum hemipteran pest control.

Different binding properties of odorant-binding protein 8 to insecticides in Orius sauteri.

Chemical sensing systems are vital in the growth and development of insects. Orius sauteri (Poppius) (Hemiptera: Anthocoridae) is an important natural enemy of many pests. The molecular mechanism of odorant binding proteins (OBPs) binding with common insecticides is still unknow in O. sauteri. In this study, we expressed in vitro OsauOBP8 and conducted fluorescence competition binding assay to investigate the function of OsauOBP8 to insecticides. The results showed that OsauOBP8 could bind with four common insecticides (phoxim, fenitrothion, chlorpyrifos, deltamethrin). Subsequently, we used molecular docking to predict and obtained candidate six amino acid residues (K4, K6, K13, R31, K49, K55) and then mutated. The result showed that three key residues (K4, K6, R31) play important role in OsauOBP8 bound to insecticides. Our study identified the key binding sites of OsauOBP8 to insecticides and help to better understand the molecular mechanism of OBPs to insecticides in O. sauteri.

Ecological drift during colonization drives within-host and between-host heterogeneity in an animal-associated symbiont.

Specialized host-microbe symbioses canonically show greater diversity than expected from simple models, both at the population level and within individual hosts. To understand how this heterogeneity arises, we utilize the squash bug, Anasa tristis, and its bacterial symbionts in the genus Caballeronia. We modulate symbiont bottleneck size and inoculum composition during colonization to demonstrate the significance of ecological drift, the noisy fluctuations in community composition due to demographic stochasticity. Consistent with predictions from the neutral theory of biodiversity, we found that ecological drift alone can account for heterogeneity in symbiont community composition between hosts, even when 2 strains are nearly genetically identical. When acting on competing strains, ecological drift can maintain symbiont genetic diversity among different hosts by stochastically determining the dominant strain within each host. Finally, ecological drift mediates heterogeneity in isogenic symbiont populations even within a single host, along a consistent gradient running the anterior-posterior axis of the symbiotic organ. Our results demonstrate that symbiont population structure across scales does not necessarily require host-mediated selection, as it can emerge as a result of ecological drift acting on both isogenic and unrelated competitors. Our findings illuminate the processes that might affect symbiont transmission, coinfection, and population structure in nature, which can drive the evolution of host-microbe symbioses and microbe-microbe interactions within host-associated microbiomes.

Identifying a potentially invasive population in the native range of a species: The enlightenment from the phylogeography of the yellow spotted stink bug, Erthesina fullo (Hemiptera: Pentatomidae).

The yellow spotted stink bug (YSSB), Erthesina fullo (Thunberg, 1783) is an important Asian pest that has recently successfully invaded Europe and an excellent material for research on the initial stage of biological invasion. Here, we reported the native evolutionary history, recent invasion history, and potential invasion threats of YSSB for the first time based on population genetic methods [using double digest restriction-site associated DNA (ddRAD) data and mitochondrial COI and CYTB] and ecological niche modelling. The results showed that four lineages (east, west, southwest, and Hainan Island) were established in the native range with a strong east-west differentiation phylogeographical structure, and the violent climate fluctuation might cause population divergence during the Middle and Upper Pleistocene. In addition, land bridges and monsoon promote dispersal and directional genetic exchanging between island populations and neighboring continental populations. The east lineage (EA) was identified as the source of invasion in Albania. EA had the widest geographical distribution among all other lineages, with a star-like haplotype network with the main haplotype as the core. It also had a rapid population expansion history, indicating that the source lineage might have stronger diffusion ability and adaptability. Our findings provided a significant biological basis for fine tracking of invasive source at the lineage or population level and promote early invasion warning of potential invasive species on a much subtler lineage level.

Foveavelia, a new South American genus of Veliinae (Hemiptera: Heteroptera: Veliidae).

Background: Semiaquatic bugs (Hemiptera: Heteroptera: Gerromorpha) are distributed worldwide and play fundamental roles in limnic ecosystems. They are the most successful group of organisms to occupy the air-water interface, are important models to study ecology and evolution, and can be relevant tools in biomonitoring. Veliidae is the second most speciose family of semiaquatic bugs, but its internal classification, including subfamilies and genera, is artificial and based on symplesiomorphies. One of these non-monophyletic entities is Paravelia Breddin, 1898, the largest genus in the subfamily Veliinae. Results: In an effort to better classify the Veliinae, we describe Foveavelia to hold five South American species previously placed in Paravelia. The new genus is characterized by the following combination of features: unusual coarse cuticular punctures throughout the thorax and abdomen; a pair of small, frosty, pubescent areas formed by a very dense layer of short setae on the anterior lobe of the pronotum; fore tibial grasping comb present only in males; middle tibia with a row of elongate dark-brown trichobothria-like setae on the distal third, decreasing in size distally; macropterous specimens with the apical macula of the forewings elongate and constricted at mid-length, reaching the wing apex; and the male proctiger with a pair of anterodorsal projections. Besides the description, a key to the species of Foveavelia is provided, accompanied by illustrations and a species distribution map.

Ethno-entomotherapeutic and metabolite profiling of Coridius chinensis (Dallas), a traditional edible insect species of North-East India.

Edible insects possess several health enhancing properties and play an important role in human nutrition. Coridius chinensis is an edible insect that is considered food and claimed as traditional medicine. In the present study, nutritional contents, chemical composition, antioxidant, and anti-inflammatory properties of C. chinensis were analyzed. It was found that the insect sample contains 50.46% moisture, 44.65% protein, 4.45% carbohydrate, 39.42% crude fats, 3.53% ash and 576.11 (Kcal/100 g) energy. Our study highlighted the presence of a significant amount of phenol and flavonoids. The C. chinensis hydro-alcoholic extract showed high antioxidant property and anti-inflammatory activity. GCMS analysis identified 61 volatile compounds. LC-MS analysis of hydroalcoholic extract of C. chinensis revealed the presence of compounds such as etodolac glucuronide, morphine 3-glucuronide, ecgonine, ecgonine methyl ester, sufentanil, and palmitoyl ethanololamide. These findings suggest that C. chinensis species can be employed as a valuable food source with excellent therapeutic properties.

Corythauma ayyari (Insecta, Heteroptera, Tingidae) depends on its host plant to spread in Europe.

Biological invasions increase with the intensity of globalization, human activities, and climate change. Insects represent a high potential of invasive species due to their adaptability to new environment. We analysed here the potential of an Asian phytophagous bug, Corythauma ayyari (Heteroptera, Tingidae) to become widespread, recently recorded in Europe, and that depends on Jasminum spp., an ornamental plant widespread in Europe. We modelled its current distribution, projected it into the future and tested its niche overlap between native and invaded areas. When considering the host plants as environmental variables, the analysis shows that C. ayyari shifted to a new ecological niche but its distribution is restricted by its host plant distribution. Including or excluding the host plants as environmental variables has an impact on C. ayyari distribution. We recommend to consider host plant interactions when dealing with niche modelling of phytophagous species.

Shared phylogeographic patterns and environmental responses of co-distributed soybean pests: Insights from comparative phylogeographic studies of Riptortus pedestris and Riptortus linearis in the subtropics of East Asia.

Comparative phylogeographic studies of closely related species sharing co-distribution areas can elucidate the role of shared historical factors and environmental changes in shaping their phylogeographic pattern. The bean bugs, Riptortus pedestris and Riptortus linearis, which both inhabit subtropical regions in East Asia, are recognized as highly destructive soybean pests. Many previous studies have investigated the biological characteristics, pheromones, chemicals and control mechanisms of these two pests, but few studies have explored their phylogeographic patterns and underlying factors. In this study, we generated a double-digest restriction site-associated DNA sequencing (ddRAD-seq) dataset to investigate phylogeographic patterns and construct ecological niche models (ENM) for both Riptortus species. Our findings revealed similar niche occupancies and population genetic structures between the two species, with each comprising two phylogeographic lineages (i.e., the mainland China and the Indochina Peninsula clades) that diverged approximately 0.1 and 0.3 million years ago, respectively. This divergence likely resulted from the combined effects of temperatures variation and geographical barriers in the mountainous regions of Southwest China. Further demographic history and ENM analyses suggested that both pests underwent rapid expansion prior to the Last Glacial Maximum (LGM). Furthermore, ENM predicts a northward shift of both pests into new soybean-producing regions due to global warming. Our study indicated that co-distribution soybean pests with overlapping ecological niches and similar life histories in subtropical regions of East Asia exhibit congruent phylogeographic and demographic patterns in response to shared historical biogeographic drivers.

Checklist and distribution of Nepomorpha (Hemiptera: Heteroptera) from China.

An updated catalog of the infraorder Nepomorpha from China is provided based on literature reports, museum specimens, and field collections. In total, 214 species of Nepomorpha are listed in 6 superfamilies, 11 families, and 37 genera, including: Aphelocheiridae (1 genus, 27 species), Belostomatidae (3 genera, 7 species), Corixidae (9 genera, 52 species), Gelastocoridae (1 genus, 3 species), Helotrephidae (5 genera, 25 species), Micronectidae (1 genus, 28 species), Naucoridae (7 genera, 12 species), Nepidae (5 genera, 21 species), Notonectidae (4 genera, 32 species), Ochteridae (1 genus, 2 species) and Pleidae (1 genus, 5 species). Paraplea liturata (Fieber, 1844) is reported from mainland China for the first time. Distribution maps are provided for most species and are based on museum specimens and our field collections.

Nanoparticle-delivered RNAi-based pesticide target screening for the rice pest white-backed planthopper and risk assessment for a natural predator.

Vacuolar-type (H+)-ATPase (vATPase) is a conserved multi-subunit eukaryotic enzyme composed of 14 subunits that form a functional complex consisting of an ATP-hydrolytic domain (V1) and a proton-translocation domain (V0). ATP hydrolysis and subsequent H+ translocation rely heavily on a fully assembled V1/V0 complex. Since vATPase is crucial for insect survival, it is a viable molecular target for pest control. However, detailed functional analyses of the 14 subunits and their suitability for pest control have not been fully explored in a single insect species. In this study, we identified 22 vATPase subunit transcripts that correspond to 13 subunits (A1, A2, B, C, D, E, F, G, H, a1, a2, c and d) in the white-backed planthopper (WBPH), Sogatella furcifera, a major hemipteran pest of rice. RNAi screens using microinjection and spray-based methods revealed that the SfVHA-F, SfVHA-a2 and SfVHA-c2 subunits are critical. Furthermore, star polymer (SPc) nanoparticles were utilized to conduct spray-induced and nanoparticle-delivered gene silencing (SI-NDGS) to evaluate the pest control efficacy of RNAi targeting the SfVHA-F, SfVHA-a2 and SfVHA-c2 transcripts. Target mRNA levels and vATPase enzymatic activity were both reduced. Honeydew excreta was likewise reduced in WBPH treated with dsRNAs targeting SfVHA-F, SfVHA-a2 and SfVHA-c2. To assess the environmental safety of the nanoparticle-wrapped dsRNAs, Cyrtorhinus lividipennis Reuter, a major natural enemy of planthoppers, was also sprayed with dsRNAs targeting SfVHA-F, SfVHA-a2 and SfVHA-c2. Post-spray effects of dsSfVHA-a2 and dsSfVHA-c2 on C. lividipennis were innocuous. This study identifies SfVHA-a2 and SfVHA-c2 as promising targets for biorational control of WBPH and lays the foundation for developing environment-friendly RNAi biopesticides.

Leveraging satellite observations to reveal ecological drivers of pest densities across landscapes.

Landscape ecologists have long suggested that pest abundances increase in simplified, monoculture landscapes. However, tests of this theory often fail to predict pest population sizes in real-world agricultural fields. These failures may arise not only from variation in pest ecology, but also from the widespread use of categorical land-use maps that do not adequately characterize habitat-availability for pests. We used 1163 field-year observations of Lygus hesperus (Western Tarnished Plant Bug) densities in California cotton fields to determine whether integrating remotely-sensed metrics of vegetation productivity and phenology into pest models could improve pest abundance analysis and prediction. Because L. hesperus often overwinters in non-crop vegetation, we predicted that pest abundances would peak on farms surrounded by more non-crop vegetation, especially when the non-crop vegetation is initially productive but then dries down early in the year, causing the pest to disperse into cotton fields. We found that the effect of non-crop habitat on pest densities varied across latitudes, with a positive relationship in the north and a negative one in the south. Aligning with our hypotheses, models predicted that L. hesperus densities were 35 times higher on farms surrounded by high versus low productivity non-crop vegetation (EVI area 350 vs. 50) and 2.8 times higher when dormancy occurred earlier versus later in the year (May 15 vs. June 30). Despite these strong and significant effects, we found that integrating these remote-sensing variables into land-use models only marginally improved pest density predictions in cotton compared to models with categorical land cover metrics alone. Together, our work suggests that the remote sensing variables analyzed here can advance our understanding of pest ecology, but not yet substantively increase the accuracy of pest abundance predictions.

Describe the morphology and mitochondrial genome of Mecidea indica Dallas, 1851 (Hemiptera, Pentatomidae), with its phylogenetic position.

We here describe the external morphology and complete mitochondrial genome characteristics of Mecidea indica Dallas, 1851, and clarify the evolutionary rate and divergence time. The M. indica mitochondrial genome length is 15,670 bp, and it exhibits a typical high A+T-skew (76.31%). The sequence shows strong synteny with the original gene arrangement of Drosophila yakuba Burla, 1954 without rearrangement. The M. indica mitochondrial genome characteristics were analyzed, and phylogenetic trees of Pentatomidae were reconstructed using Bayesian methods based on different datasets of the mitochondrial genome datasets. Phylogenetic analysis shows that M. indica belongs to Pentaotominae and form a sister-group with Anaxilaus musgravei Gross, 1976, and Asopinae is highly supported as monophyletic. Molecular clock analysis estimates a divergence time of Pentatomidae of 122.75 Mya (95% HPD: 98.76-145.43 Mya), within the Mesozoic Cretaceous; the divergence time of M. indica and A. musgravii was no later than 50.50 Mya (95% HPD: 37.20-64.80 Mya). In addition, the divergence time of Asopinae was 62.32 Mya (95% HPD: 47.08-78.23 Mya), which was in the Paleogene of the Cenozoic era. This study is of great significance for reconstructing the phylogeny of Pentatomidae and providing insights into its evolutionary history.

Salivary proteins potentially derived from horizontal gene transfer are critical for salivary sheath formation and other feeding processes.

Herbivorous insects employ an array of salivary proteins to aid feeding. However, the mechanisms behind the recruitment and evolution of these genes to mediate plant-insect interactions remain poorly understood. Here, we report a potential horizontal gene transfer (HGT) event from bacteria to an ancestral bug of Eutrichophora. The acquired genes subsequently underwent duplications and evolved through co-option. We annotated them as horizontal-transferred, Eutrichophora-specific salivary protein (HESPs) according to their origin and function. In Riptortus pedestris (Coreoidea), all nine HESPs are secreted into plants during feeding. The RpHESP4 to RpHESP8 are recently duplicated and found to be indispensable for salivary sheath formation. Silencing of RpHESP4-8 increases the difficulty of R. pedestris in probing the soybean, and the treated insects display a decreased survivability. Although silencing the other RpHESPs does not affect the salivary sheath formation, negative effects are also observed. In Pyrrhocoris apterus (Pyrrhocoroidea), five out of six PaHESPs are secretory salivary proteins, with PaHESP3 being critical for insect survival. The PaHESP5, while important for insects, no longer functions as a salivary protein. Our results provide insight into the potential origin of insect saliva and shed light on the evolution of salivary proteins.

Effects of photoperiod and temperature on the developmental duration and diapause in Dolycoris baccarum (Heteroptera: Pentatomidae) from Hohhot, Inner Mongolia, China.

The shield bug, Dolycoris baccarum (L.) (Heteroptera: Pentatomidae), is widely distributed across Asia and Europe. At high latitudes, it overwinters, as adult in diapause, which then becomes the insect source for the following year. To fully understand the developmental duration and diapause characteristics of D. baccarum, the effects of photoperiod and temperature were studied in a population from Hohhot, Inner Mongolia, China. The results indicated that the developmental duration was significantly prolonged at temperatures of 20 or 25 °C, with a prolonged light period; however, when the light period was prolonged to 16L:8D and 18L:6D, the developmental duration was shortened significantly. Furthermore, the developmental duration was also shortened significantly with increasing temperature, when the photoperiod was 12L:12D for short days and 16L:8D for long days. All individuals entered diapause under short-day conditions of 10L:14D and 12L:12D at a temperature of 20 °C; however, the diapause rate decreased significantly under 14L:10D and 16L:8D photoperiods, and the diapause rate decreased significantly at a temperature of 25 °C with prolonged photoperiod. Interestingly, when the photoperiod was fixed at 12L:12D, the diapause rates at different temperatures (20, 25, 28, and 30 °C) exceeded 95%; while the effect of temperature on diapauses was nonsignificant under this photoperiod, it was still sensitive to the photoperiod; at a photoperiod of 16L:8D, the effect of temperature on the diapause rate was noticeable, and the diapause rate decreased significantly with increasing temperature.