Fact-checked by Grok 2 weeks ago

Hemiptera

Hemiptera is an order of insects commonly known as true bugs, comprising more than 100,000 described species worldwide and characterized by their piercing-sucking mouthparts formed into a tubular proboscis (rostrum) used to extract fluids from plants, animals, or other sources.^[1]^[2] This diverse group undergoes incomplete metamorphosis, with eggs hatching into nymphs that resemble smaller, wingless versions of the adults and develop through several instars before molting into winged adults.^[3] Hemipterans are found in nearly all terrestrial and freshwater aquatic habitats globally, from forests and fields to ponds and streams, and include both free-living and sedentary forms.^[1]^[2] The order Hemiptera is divided into three main suborders: Sternorrhyncha, which includes aphids, whiteflies, scale insects, and mealybugs that primarily feed on plant sap and often form mutualistic relationships with ants; Auchenorrhyncha, encompassing cicadas, leafhoppers, planthoppers, and treehoppers known for their sound-producing abilities and jumping locomotion; and Heteroptera, the true bugs featuring hemelytra (forewings with a leathery basal portion and membranous apical region) and a wide range of feeding habits from herbivory to predation and bloodsucking. A fourth, smaller suborder, Coleorrhyncha, includes bark lice-like insects.^[1]^[2] With more than 42,000 species in Heteroptera alone, the order exhibits remarkable morphological variation, including body sizes from under 1 mm to over 10 cm, and antennae that are typically long and segmented.^[2]^[3] Ecologically, Hemiptera play pivotal roles as herbivores that can damage crops by sucking plant juices and transmitting viruses, such as aphids vectoring cucumber mosaic virus, while predatory species like assassin bugs control pest populations.^[1] Aquatic and semi-aquatic forms, numbering around 4,000 species, inhabit freshwater ecosystems where they prey on smaller invertebrates like mosquito larvae or scavenge detritus, serving as bioindicators of water quality and food for fish and amphibians.^[1]^[4] Economically, certain Hemiptera are significant pests in agriculture and human health—bed bugs and kissing bugs transmit diseases like Chagas—yet others, such as cochineal scales, provide natural dyes, and predatory bugs aid in biological pest control.^[2]^[4]^[5]

Taxonomy and phylogeny

Classification

The order Hemiptera is divided into four suborders: Sternorrhyncha, Coleorrhyncha, Auchenorrhyncha, and Heteroptera.^[5] Sternorrhyncha comprises primarily small, soft-bodied insects that feed on plant sap, including aphids, whiteflies, psyllids, and scale insects, and is further divided into superfamilies such as Aphidoidea and Coccoidea.^[5] Coleorrhyncha includes a single extant family, Peloridiidae (about 40 species), known as moss bugs, which are Gondwanan relicts with archaic features and feed on mosses and liverworts. Auchenorrhyncha includes hoppers, planthoppers, treehoppers, and cicadas, characterized by their ability to produce sound and jump, with two main infraorders: Fulgoromorpha (planthoppers and allies) and Cicadomorpha (cicadas and leafhoppers), the latter encompassing superfamilies like Cicadoidea.^[6] Heteroptera, the true bugs, exhibit diverse feeding habits ranging from predation to phytophagy and even blood-feeding, organized into infraorders such as Cimicomorpha and Pentatomomorpha, with key superfamilies including Pentatomoidea (shield bugs and allies) and Reduvioidea (assassin bugs).^[5] Major families within Hemiptera highlight the order's diversity. The Aphididae (aphids), in Sternorrhyncha: Aphidoidea, contains approximately 5,000 species, known for their role as plant pests and virus vectors.^[7] The Cicadidae (cicadas), in Auchenorrhyncha: Cicadomorpha: Cicadoidea, includes over 3,000 species, notable for their periodic emergences and loud calls produced by males.^[8] In Heteroptera: Pentatomomorpha: Pentatomoidea, the Pentatomidae (stink bugs) comprises about 4,700 species, many of which are herbivorous and release defensive odors when disturbed.^[9] The Reduviidae (assassin bugs), in Heteroptera: Cimicomorpha: Reduvioidea, encompasses roughly 7,000 species, primarily predatory insects that use a proboscis to inject venom into prey.^[10] Recent taxonomic updates have refined Hemiptera classification. In 2024, a phylogenomic and morphological analysis revised the Reduviidae, reducing the number of subfamilies from 25 to 19 and establishing 40 tribes, while describing three new subfamilies to better reflect evolutionary relationships.^[11] Nomenclatural changes in Auchenorrhyncha, proposed in 2020 and influencing ongoing revisions into 2025, addressed homonyms by introducing replacement names for several genera and higher taxa across the suborder.^[12] Additionally, in 2023, a new family was described from Chinese taxa, contributing to the recognition of novel lineages within Hemiptera based on specimens from the region.^[13]

Phylogenetic relationships

Hemiptera belongs to the hemipteroid insects within the Paraneoptera superorder of Insecta, encompassing over 100,000 described species characterized by their hemimetabolous development and specialized feeding adaptations. Phylogenomic analyses using transcriptomes and genomes from 193 hemipteroid species strongly support Thysanoptera (thrips) as the sister group to Hemiptera, forming the monophyletic Condylognatha clade; this relationship is backed by 100% bootstrap support from 2,395 orthologous genes and an estimated divergence around 407 million years ago in the Devonian period.^[14] The broader Hemipterida clade includes Psocodea as an outgroup to Condylognatha, with monophyly of all three hemipteroid orders (Psocodea, Thysanoptera, Hemiptera) affirmed by quartet-based likelihood mapping across 859,518 aligned amino acid positions.^[14] The internal phylogeny of Hemiptera is monophyletic, unified by synapomorphies such as the elongate, piercing-sucking rostrum for fluid ingestion and the presence of a buccula (a lateral groove on the head). Among the suborders, Sternorrhyncha is positioned as basal, sister to all remaining hemipterans, followed by Coleorrhyncha as sister to the monophyletic Auchenorrhyncha, with Heteroptera emerging as the derived lineage; this topology reflects a progression from plant sap-feeding in basal groups to diverse predatory and phytophagous habits in Heteroptera.^[15] Auchenorrhyncha exhibits internal structure with Cicadoidea + Cercopoidea sister to Membracoidea, while Heteroptera's monophyly is marked by additional synapomorphies like scent glands and forewing differentiation into coriaceous basal and membranous distal regions.^[15] Molecular evidence has solidified these relationships through comprehensive genomic and transcriptomic datasets. A 2024 study sequenced genomes from 64 Hemiptera species spanning 15 superfamilies (including 13 from Sternorrhyncha, 30 from Auchenorrhyncha, and 21 from Heteroptera) and integrated transcriptomes from two additional scale insects, analyzing 1,625 nuclear loci across 315 species via Bayesian inference, maximum likelihood, and four-cluster likelihood mapping.^[15] These methods confirmed Sternorrhyncha's basal position, with Aphididae (e.g., species like Acyrthosiphon pisum) robustly nested within Aphidoidea as sister to Phylloxeroidea; Bayesian analyses further delineated divergences, such as the split between Auchenorrhyncha and Heteroptera around 250–300 million years ago.^[15] Historical controversies in Hemiptera phylogeny included debates over Heteroptera monophyly, with some early morphological studies suggesting paraphyly due to convergent traits in feeding and wing venation, but these have been resolved by molecular data. Recent transcriptomic analyses, including those from the 2024 dataset, provide unequivocal support for Heteroptera as a clade, emphasizing its predatory ancestry and subsequent phytophagous radiations, while also clarifying longstanding uncertainties in Auchenorrhyncha relationships.^[15]^[14]

Fossil record and evolutionary history

The fossil record of Hemiptera extends back to the Late Carboniferous, with the oldest known specimens dating to approximately 320 million years ago (Ma), including early lineages such as Protoprosbolidae and Aviorrhynchidae.^[16] Bayesian analyses of the fossil record estimate the origin of crown-group Hemiptera in the Bashkirian stage of the early Pennsylvanian, around 321 Ma (95% credible interval: 311–344 Ma), suggesting stem-hemipterans appeared shortly after the diversification of early polyneopteran insects.^[17] These initial fossils, primarily from European deposits like the Commentry Basin in France, represent primitive forms with basic piercing-sucking mouthparts adapted to gymnosperm hosts, predating the Permian diversification of more derived taxa.^[17] Diversification of Hemiptera accelerated during the Permian, particularly in the Guadalupian and Lopingian stages, where birth-death models indicate a significant radiation involving at least 27 new families, such as Prosbolomorpha and early Psyllaeformia, coinciding with expanding terrestrial ecosystems.^[17] A 2025 Bayesian study using birth-death shift models on 11,840 fossil occurrences revealed further peaks in net diversification during the Early Jurassic (origination rate ~0.13) and Early Cretaceous (~117–102 Ma, with up to 100 genera), driven by ecological opportunities in angiosperm-dominated habitats.^[16] Within suborders, Sternorrhyncha exhibited a pronounced radiation in the mid-Cretaceous (Aptian–Cenomanian), linked to the rise of flowering plants, as evidenced by amber fossils showing shifts from gymnosperm to angiosperm feeding; for instance, early Aleyrodomorpha and Psyllodea lineages diversified rapidly, with 19 of 30 extant planthopper families emerging by the Late Cretaceous.^[18] Heteroptera, in contrast, peaked in the Early Jurassic and Late Jurassic–Early Cretaceous, reflecting adaptations to predatory and semi-aquatic niches.^[16] Major extinction events profoundly shaped Hemiptera evolution, with the end-Permian mass extinction (~252 Ma) causing the highest recorded extinction rate (~0.6 per million years) and turnover of ancient lineages like Ignotalidae and Pereboriidae, though some clades survived into the Triassic.^[16] The Cretaceous–Paleogene (K-Pg) boundary (~66 Ma) contributed to a diversity decline in the early Paleogene, particularly affecting specialized groups, but was followed by Cenozoic recoveries, including radiations around 36 Ma and 17 Ma that established modern faunas.^[16] Post-K-Pg patterns show correlated origination with angiosperm expansion (correlation coefficient ω ≈ 0.78) and extinctions tied to declines in non-polypod ferns (ω ≈ 0.8), underscoring plant-insect co-evolutionary dynamics.^[16] Key fossil taxa illustrate these patterns, such as the Carboniferous Protoprosbole straeleni (~315–325 Ma), an early hemipteran with elongate proboscis, and Permian Curvicubitidae, which highlight pre-extinction diversity in Pereborioidea.^[17] Triassic records include early Heteroptera like Hylicellidae and Ipsviciidae from the Norian Cow Branch Formation (~220 Ma), marking the suborder's emergence with advanced scent glands.^[19] Eocene amber deposits, particularly from Baltic and French sources (~45–50 Ma), preserve diverse inclusions like the first fossil Dipsocoridae (e.g., Oedancala gallica), revealing minute, ground-dwelling heteropterans, and Sternorrhyncha such as Dingla shagria from earlier Cretaceous ambers, which document trophic interactions in resin-trapped ecosystems.^[20]^[18] These amber fossils provide exceptional detail on wing venation and body preservation, bridging gaps in the compression fossil record.^[18]

Diversity

Species richness and distribution

Hemiptera, the order of true bugs, encompasses approximately 100,000 described species worldwide as of 2025, making it the fifth most species-rich insect order after Coleoptera, Lepidoptera, Diptera, and Hymenoptera.^[21] Estimates suggest the true total exceeds 200,000 species, accounting for undescribed diversity, particularly in tropical regions.^[22] The order is divided into three major suborders: Sternorrhyncha with around 18,000 described species, primarily comprising aphids, scale insects, whiteflies, and psyllids; Auchenorrhyncha with over 42,000 species, including cicadas, leafhoppers, and planthoppers; and Heteroptera with more than 45,000 species, encompassing predatory and herbivorous true bugs.^[23]^[24]^[25] Hemiptera exhibit a cosmopolitan distribution, occurring on every continent except Antarctica, with the highest species richness concentrated in tropical and subtropical regions.^[26] Biodiversity hotspots include Southeast Asia and the Amazon Basin, where environmental heterogeneity supports elevated diversity, particularly among phytophagous and aquatic taxa.^[27] Approximately 4,000 species are aquatic or semiaquatic, predominantly in the Heteroptera suborder, and are distributed worldwide in freshwater habitats such as streams, ponds, and wetlands.^[1] Patterns of endemism vary by suborder and region, with high levels in isolated tropical areas, while Hemiptera demonstrate notable invasive potential due to traits like parthenogenesis and broad host ranges, as evidenced by a 2024 global analysis of over 1,600 non-native establishments.^[21] Recent taxonomic efforts have added significantly to known diversity, including 252 new species described from China between 2022 and 2023, highlighting ongoing discoveries in megadiverse areas.^[28]

Major groups and examples

The suborder Sternorrhyncha encompasses plant-sap feeding insects such as aphids and scale insects, notable for their sessile or colonial lifestyles and ability to reproduce parthenogenetically. The family Aphididae includes the green peach aphid (Myzus persicae), a cosmopolitan pest that colonizes a wide range of crops like vegetables and ornamentals, producing honeydew that promotes sooty mold growth.^[29] In the superfamily Coccoidea, the family Diaspididae features the San Jose scale (Quadraspidiotus perniciosus), an invasive armored scale that forms waxy coverings and infests fruit trees, weakening hosts through phloem feeding.^[30] The suborder Auchenorrhyncha comprises hopping insects like cicadas and planthoppers, distinguished by their robust hind legs adapted for jumping and, in some cases, loud acoustic signaling. The family Cicadidae is represented by the periodical cicadas of the genus Magicicada, which emerge en masse after 13 or 17 years underground, engaging in synchronized chorusing to attract mates and overwhelm predators.^[31] The family Delphacidae includes various planthoppers, such as the brown planthopper (Nilaparvata lugens), which uses stealthy feeding on rice stems and can produce frothy spittle for protection during nymphal stages.^[31] The suborder Heteroptera, or true bugs, exhibits diverse habits from predation to herbivory, often characterized by defensive chemical secretions. The family Pentatomidae includes the brown marmorated stink bug (Halyomorpha halys), an invasive species from Asia that aggregates on crops and structures, releasing a pungent odor when disturbed.^[32] In aquatic habitats, the family Nepidae features water scorpions like Ranatra species, which ambush prey using raptorial front legs and breathe through a tail-like siphon.

Morphology

General body plan

Hemiptera exhibit the typical insect body plan, divided into three primary regions: the head, thorax, and abdomen. The head is generally small and triangular, bearing compound eyes, ocelli in some species, and antennae that vary from short and concealed to long and segmented. The thorax is compact, consisting of three segments (pro-, meso-, and metathorax) that support three pairs of legs and, in winged forms, two pairs of wings. The abdomen is elongate and segmented, typically comprising 10-11 visible segments in adults, enclosing the digestive, reproductive, and respiratory systems. This tripartite structure facilitates diverse adaptations for feeding, locomotion, and reproduction across the order.^[33]^[1] Size variation among Hemiptera is extensive, ranging from approximately 0.4 mm in some tiny aphids of the family Aphididae to up to 12 cm in large aquatic species such as giant water bugs (Lethocerus spp.). This disparity reflects their ecological diversity, with minute forms often colonizing dense plant tissues and larger ones inhabiting aquatic environments.^[34]^[35] Coloration in Hemiptera is diverse and often cryptic, featuring mottled greens, browns, or grays that blend with vegetation or substrates for camouflage against predators. Many species produce waxy secretions or froth to enhance concealment and prevent desiccation. Sexual dimorphism occurs in several groups, notably in aphids and scale insects, where females are frequently wingless and apterous while males develop wings for dispersal.^[36]^[37] The exoskeleton of Hemiptera is composed of chitin, forming a rigid yet lightweight cuticle that protects internal organs and allows flexibility at segmental joints. In the suborder Heteroptera, specialized scent glands are embedded in the thorax and abdomen, opening externally to release volatile defensive compounds when threatened. In Heteroptera, the forewings are characteristically hemelytra, with a leathery corium basally and a translucent membrane apically, contributing to the overall dorsally flattened profile.^[1]^[38]

Mouthparts and feeding structures

Hemiptera are characterized by specialized piercing-sucking mouthparts that form a prominent rostrum, consisting of a segmented labium that serves as a protective sheath for four elongated stylets derived from the modified mandibles and maxillae.^[39] The two mandibular stylets are typically outer and rigid, functioning to puncture host tissues, while the two maxillary stylets interlock to create a central food canal for ingesting liquids and a parallel salivary canal for injecting secretions, with the canals often connected by interlocking structures for stability during penetration.^[40] This arrangement allows precise insertion into vascular tissues or soft body parts, distinguishing Hemiptera from other insect orders with chewing or lapping mouthparts. Variations in rostrum and stylet morphology reflect dietary adaptations across Hemiptera subgroups. In herbivorous species like aphids (Sternorrhyncha), the rostrum is relatively short and attached ventrally near the head's rear, suited for accessing phloem; however, in genera such as Stomaphis, the stylets can exceed 10 mm in length, enabling penetration of thick tree bark, with the ability to invert and coil within a specialized pouch called the crumena during non-feeding states.^[41] Predatory and parasitic forms, such as in Reduviidae (Heteroptera), exhibit longer, more robust rostra with asymmetrical stylets featuring barbs or ridges for anchoring in prey, as seen in Haematoloecha nigrorufa where mandibular stylets have spatulate tips with transverse ridges for breaching millipede cuticles.^[42] In Auchenorrhyncha, the rostrum often coils when retracted and attaches closer to the coxae, facilitating diverse plant-feeding strategies.^[39] Salivary secretions in Hemiptera are delivered through the maxillary stylet's salivary canal and contain specialized enzymes tailored to feeding ecology. Herbivores like aphids produce cellulases in their saliva to break down plant cell walls, aiding tissue penetration and nutrient extraction, as first documented in the Russian wheat aphid Diuraphis noxia.^[43] In hematophagous species, such as triatomine bugs (Reduviidae), saliva includes anticoagulants like serine protease inhibitors to prevent blood clotting during feeding, ensuring uninterrupted flow from host vessels.^[44] The piercing-sucking mouthparts represent a key synapomorphy defining the order Hemiptera, marking an evolutionary transition from the ancestral mandibulate (chewing) condition in Polyneoptera to a haustellate system optimized for fluid diets.^[45] This innovation likely originated in the early Permian, coinciding with the diversification of hemipteroids and a shift toward phytophagy or predation on fluids, as evidenced by fossil mouthpart impressions from the Carboniferous.^[46]

Wing structure and polymorphism

The wings of Hemiptera exhibit considerable diversity across suborders, reflecting adaptations to various ecological niches. In the suborder Heteroptera, the forewings, known as hemelytra, are characteristically divided into a coriaceous (leathery or hardened) basal portion and a membranous apical portion, providing both protection and flexibility.^[47]^[48] The hindwings are entirely membranous and fold beneath the hemelytra when at rest.^[49] In contrast, members of the suborder Auchenorrhyncha possess forewings termed tegmina, which are typically leathery or parchment-like and extend over the abdomen, with hindwings that are membranous and often held in a fan-like position under the tegmina.^[50]^[51] Wing polymorphism is a prominent feature in many Hemiptera, particularly in response to environmental pressures, allowing individuals to trade off flight capability for other fitness advantages such as enhanced reproduction or energy conservation. Common morphs include macropterous (fully winged), brachypterous (short-winged), and apterous (wingless) forms, with the latter two often occurring in stable or crowded conditions.^[52]^[53] In aphids (Sternorrhyncha), this polymorphism is environmentally induced; for instance, high population density, deteriorating host plant quality, or predation cues trigger the development of winged morphs for dispersal, while favorable conditions favor wingless forms that allocate resources to rapid reproduction.^[54]^[55] Such phenotypic plasticity enables aphids to colonize new habitats during seasonal migrations.^[56] Venation patterns in Hemipteran wings vary significantly among suborders and contribute to structural integrity and functional roles. In Sternorrhyncha, such as aphids and scale insects, wing venation is highly reduced, often consisting of only a few prominent veins with minimal branching, which correlates with the small size and limited flight demands of these groups.^[57]^[58] Conversely, in Auchenorrhyncha and Heteroptera, venation is more complex and reticulate, featuring numerous cross-veins that enhance wing rigidity and facilitate resonance properties.^[59]^[60] Unlike the order Diptera, where the hindwings are modified into halteres for balance during flight, Hemiptera possess two pairs of fully developed wings without such reductions or specialized sensory structures.^[61]^[47] This configuration supports direct flight mechanics in winged forms.^[62]

Physiology and life history

Sensory systems and communication

Hemiptera exhibit a diverse array of sensory systems adapted to their ecological niches, including vision, olfaction, and mechanoreception, which facilitate foraging, host location, and conspecific interactions. The visual system primarily consists of compound eyes, with three ocelli present in most adult Heteroptera for detecting light intensity and motion, though ocelli are typically absent in adult Sternorrhyncha such as aphids.^[63] Compound eyes in Hemiptera provide panoramic vision and color perception, enabling the detection of environmental cues like plant silhouettes and contrasts. In aphids, for example, the compound eyes support trichromatic vision with photoreceptor sensitivities peaking in the ultraviolet (around 350 nm), blue-green (around 450-500 nm), and green (around 520-530 nm) spectra, allowing discrimination of host plants from non-host backgrounds through UV-reflective properties of foliage.^[64] This UV sensitivity is crucial for host finding, as aphids preferentially orient toward yellow-green targets that mimic plant colors under natural skylight, which includes significant UV components, while avoiding high-UV reflecting surfaces.^[65] Olfactory and chemosensory capabilities in Hemiptera are mediated primarily by antennae, which display morphological variations across suborders. In Sternorrhyncha, antennae are often filiform or flagellate, consisting of a short scape, pedicel, and multi-segmented flagellum equipped with sensilla for detecting plant volatiles and pheromones; for instance, psyllids exhibit numerous olfactory sensilla on their filiform antennae to locate host plants.^[66] In contrast, Heteroptera antennae are typically segmented into four or five parts, with elaborate trichoid and basiconic sensilla concentrated on the distal segments for enhanced pheromone detection, enabling species-specific mate recognition and aggregation.^[67] These antennal sensilla house olfactory receptor neurons that respond to sex and alarm pheromones, such as the aggregation pheromones in stink bugs (Pentatomidae), which guide conspecifics to food sources or mating sites.^[67] Mechanoreceptors, particularly hair-like sensilla (trichoid sensilla), are distributed across the body and appendages in Hemiptera, serving to detect tactile stimuli, air currents, and substrate vibrations. These sensilla feature a flexible socket and dendritic processes that transduce mechanical deflections into neural signals, with sensitivity to vibrations in the 10-1000 Hz range relevant for close-range communication.^[63] In mating contexts, such mechanoreceptors on the antennae and legs allow individuals to perceive substrate-borne vibrations produced by potential partners, facilitating pair formation without relying on airborne cues; for example, in leafhoppers (Cicadellidae), antennal trichoid sensilla help males locate calling females through vibratory signals transmitted via plant stems.^[68] Communication in Hemiptera relies heavily on non-acoustic chemical and mechanical signals, with pheromones playing a central role in coordination and defense. Aphids, for instance, release alarm pheromones like (E)-β-farnesene from the cornicles, which disperse conspecifics during predation events and can form transient chemical gradients that guide avoidance behaviors.^[69] Sex pheromones in aphids, such as nepetalactone and nepetalactol isomers, are detected via antennal sensilla to attract males, sometimes involving trail-following where males orient along pheromone deposits left by females on substrates.^[70] These chemical signals integrate with mechanosensory inputs for multimodal communication, though airborne acoustic signals are also employed in some groups for longer-range interactions.^[68]

Sound production mechanisms

Hemiptera employ diverse mechanisms for sound production, primarily serving communication, mating, and defense, with variations across suborders reflecting evolutionary adaptations. The most prominent methods include tymbal buckling in larger species like cicadas and stridulation through friction in smaller forms such as leafhoppers and true bugs. These sounds often manifest as airborne calls or substrate-borne vibrations, with frequencies typically ranging from 50 Hz to 20 kHz depending on the group.^[71]^[72]^[73] In the suborder Auchenorrhyncha, cicadas (Cicadidae) produce loud, species-specific songs using specialized tymbal organs located on the dorsolateral abdomen. Each tymbal consists of a ribbed, biconvex membrane connected to a dedicated muscle; rapid contraction buckles the ribs sequentially inward, generating a series of sharp clicks that resonate to form continuous tones. Amplification occurs via adjacent air sacs in the abdomen, which act as Helmholtz resonators, boosting sound pressure levels to over 100 dB and enabling long-distance transmission; dominant frequencies often fall between 1 and 20 kHz, with individual clicks at 4-6 kHz in species like Cyclochila australasiae. This mechanism is highly efficient, converting muscle energy directly into acoustic power, and is almost exclusively used by males for mate attraction.^[71]^[74] Stridulation, a widespread mechanism across Auchenorrhyncha and Heteroptera, involves rubbing sclerotized body parts to create friction-generated vibrations. In leafhoppers (Cicadellidae), males typically stridulate by scraping file-like structures on the forewings or costal margins against plectra on the forelegs or tegulae, producing substrate-borne signals that propagate through plants for short-range communication. These vibrations feature low-frequency peaks around 100 Hz, extending to 10 kHz with harmonics, and form patterned calls for species recognition and courtship. In planthoppers (Fulgoridae and Delphacidae), similar wing-leg friction yields comparable signals, often modulated for male-female duets.^[72]^[75] Within Heteroptera, stridulation dominates, utilizing diverse organ pairs such as a plectrum on the hind femur rubbing against a stridulitrum on the forewing edge, as seen in stink bugs (Pentatomidae like Nezara viridula). Frequencies range from 1-10 kHz, producing chirps or trills for mating or alarm. Some groups, including certain Pentatomidae and burrower bugs (Cydnidae), employ simple tymbals—vibrating abdominal tergites over air-filled chambers—yielding low-frequency pulses around 50-200 Hz. Body vibration, where the abdomen or legs tremulate against the substrate, supplements these in species like Tritomegas bicolor (Cydnidae), generating 80-150 Hz signals primarily for courtship.^[73]^[76] Sound production is notably reduced or absent in the suborder Sternorrhyncha, where aphids and scale insects rarely vocalize, though some psyllids (Psyllidae) use wing buzzing or axillary stridulation to emit weak vibrational chirps around 100-500 Hz for mating, lacking the complex organs of other hemipterans. In Heteroptera and Auchenorrhyncha, production is often sex-specific, with males generating calls to attract females, while Sternorrhyncha show minimal sexual dimorphism in this trait. These variations highlight the suborder-specific evolution of vibroacoustic signaling, with tymbals evolving independently in Auchenorrhyncha and select Heteroptera lineages.^[77]^[78]^[75]

Life cycle and development

Hemiptera exhibit hemimetabolous, or incomplete, metamorphosis, characterized by three primary life stages: egg, nymph, and adult, without a distinct pupal phase. This gradual developmental process allows nymphs to progressively resemble the adult form through a series of molts, enabling direct environmental interaction from early stages.^[36]^[79] The egg stage varies in duration and structure depending on the species, often laid in clusters on host plants or substrates. Upon hatching, nymphs emerge as miniature, wingless versions of adults, sharing similar body plans, mouthparts, and behaviors but lacking full reproductive capability and functional wings. Nymphs undergo 5 to 8 instars—developmental phases between molts—during which they grow larger and develop external wing pads, particularly in the later instars, that gradually expand and become functional upon reaching adulthood. Molting involves shedding the exoskeleton to accommodate growth, a process regulated by hormones such as juvenile hormone and ecdysone, with wing morphogenesis showing enhanced gene expression in the final two instars in species like the milkweed bug Oncopeltus fasciatus.^[36]^[79]^[80] The duration of the life cycle in Hemiptera varies widely across taxa, influenced by environmental conditions and species ecology. For instance, aphids complete development rapidly, with nymphs maturing in as little as 7 to 10 days under warm temperatures, allowing multiple annual generations and population booms. In contrast, periodical cicadas of the genus Magicicada have extended subterranean nymphal periods lasting up to 17 years before emerging as adults. Temperate Hemiptera species often incorporate diapause—a hormonally mediated dormancy—to survive unfavorable conditions like winter; for example, the brown marmorated stink bug Halyomorpha halys enters diapause as adults or late-instar nymphs, enhancing cold tolerance and overwintering success through physiological adaptations such as lowered supercooling points.^[81]^[80]^[82]

Reproductive strategies

Hemiptera exhibit diverse mating behaviors, often involving chemical and acoustic signals to facilitate mate location and courtship. In many species, sex pheromones play a crucial role in attracting conspecifics over long distances, with females typically emitting these volatile compounds from glands on the abdomen or metathorax.^[67] For instance, in leafhoppers and planthoppers (Auchenorrhyncha), substrate-borne vibrations produced by wing or body movements serve as courtship songs, allowing males to signal readiness and species identity to females.^[83] In cicadas, males generate loud acoustic calls through tymbal organs, which function in pair formation by advertising territory and attracting responsive females.^[83] A notable exception occurs in the bed bug Cimex lectularius (Cimicidae), where males employ traumatic insemination, piercing the female's abdominal wall with specialized parameres to inject sperm directly into the hemocoel, bypassing the genital tract and often leading to sexual conflict over mating frequency.^[84] Oviposition strategies in Hemiptera vary with habitat and feeding ecology, ensuring egg protection and proximity to resources. Phytophagous species, such as pentatomids and coreids (Heteroptera), typically deposit eggs in clusters on the undersides of plant leaves or within tissue crevices, providing camouflage and defense against predators.^[85] Aquatic hemipterans, like those in Corixidae, lay eggs on submerged vegetation or water surfaces, often cemented with a sticky secretion.^[85] Parental care enhances survival in some groups; for example, in giant water bugs of the subfamily Lethocerinae (Belostomatidae), females oviposit on emergent vegetation above water, after which males guard the clutches by moistening, shading, and defending them from threats until hatching.^[86] In contrast, Belostomatinae females glue eggs directly to the male's dorsal surface, where he provides brooding care, including oxygenation, for the duration of development.^[86] Parthenogenesis is prevalent in certain Hemiptera, particularly aphids (Sternorrhyncha), enabling rapid asexual reproduction under favorable conditions. Cyclic parthenogenesis in species like the pea aphid Acyrthosiphon pisum alternates 10–30 viviparous asexual generations with a single sexual generation triggered by short photoperiods and low temperatures, allowing overwintering eggs to persist.^[87] This strategy confers colonization advantages by facilitating explosive population growth and dispersal via winged morphs, without the need for mates.^[87] True thelytokous parthenogenesis, producing only females, occurs in some mirids (Heteroptera), such as Campyloneura virgula, where males are rare and reproduction proceeds asexually.^[85] Viviparity, involving internal embryonic development and live birth, has evolved independently in Hemiptera, providing protection and nutritional support. In aphids, viviparous females nourish developing embryos via pseudoplacental structures connected to maternal hemolymph, resulting in telescoped generations where granddaughters form within daughters.^[87] This maternal provisioning enhances offspring survival in unstable environments. In Heteroptera, pseudoplacental viviparity occurs in ectoparasitic polyctenids like Hesperoctenes fumarius and some anthocorids such as Physopleurella, where embryos receive nutrients directly from the female's hemocoel, analogous to adenotrophic systems in other insects.^[88]

Locomotion and behavior

Flight and dispersal

Hemiptera employ synchronous flight muscles, in which each muscle contraction is directly triggered by a neural impulse, enabling controlled wing movements during locomotion.^[89] These direct flight muscles attach to the wings or associated structures, facilitating both short bursts for escape and sustained flight in larger species such as cicadas, which can maintain powered flight over moderate distances.^[89] Unlike asynchronous muscles in some other insect orders, this synchronous mechanism limits wingbeat frequencies but allows precise control, particularly in smaller hemipterans like aphids.^[90] Dispersal in Hemiptera often relies on winged (alate) morphs, especially in aphids, where environmental cues like crowding or host plant decline induce the production of these forms for migration to new habitats.^[91] Alate aphids undertake flights that can span hundreds of kilometers, frequently aided by wind currents that carry them passively over long distances, enhancing colonization of distant plant populations.^[92] This polymorphism allows rapid spread but is balanced against apterous (wingless) morphs that prioritize reproduction over mobility in stable environments.^[91] The evolution of flight capability in Hemiptera involves significant trade-offs, including high energy costs for muscle maintenance and wing development, which reduce reproductive output in winged individuals compared to wingless ones.^[93] For instance, in planthoppers, flight-capable males exhibit lower fecundity due to resource allocation toward flight structures, though brief flights can sometimes mitigate these costs by accelerating ovarian maturation.^[94] Many hemipteran species or morphs remain apterous, restricting dispersal to walking or passive transport and limiting range expansion.^[95] Additionally, some hemipterans, such as certain reduviids, engage in nocturnal flight to minimize predation risk from diurnal visual hunters, though this exposes them to bat echolocation.^[96]

Jumping and terrestrial movement

Hemiptera exhibit diverse forms of terrestrial locomotion adapted to their habitats and lifestyles, primarily through walking, running, and jumping using modified legs. In ground-dwelling species, such as those in the families Saldidae and Rhyparochromidae, cursorial legs with elongate femora and slender tibiae facilitate rapid walking and running on soil or vegetation surfaces.^[97] These adaptations allow for efficient ground navigation, though speeds remain modest compared to specialized runners in other insect orders. In contrast, sedentary groups like aphids (Aphididae) and scale insects (Coccidae) display reduced terrestrial mobility, with adults often immobile after settling on host plants to feed.^[98] Jumping represents a key specialization for terrestrial movement in many Hemiptera, particularly in suborders Auchenorrhyncha such as froghoppers (Cercopidae) and planthoppers (Delphacidae). These insects possess hind legs with enlarged femora containing fast-contracting muscles that enable powerful leaps.^[99] A distinctive catapult-like mechanism stores elastic energy in the trochanter-femur joint during slow muscle contraction, which is then released explosively by fast extensor muscles to propel the body. In froghoppers like Philaenus spumarius, this system allows jumps reaching heights of up to 700 mm—over 100 times the body length—with takeoff velocities exceeding 4 m/s achieved in under 1 ms.^[99] Synchronous depression of both hind legs ensures stable propulsion without rotation, while tarsi pierce surfaces for traction during launch.^[100] Rapid jumping often serves as an escape response to predators in terrestrial Hemiptera. Shore bugs (Saldidae), for instance, execute short, explosive leaps from the ground to evade threats, using hind leg specialization similar to that in froghoppers but scaled for their predatory lifestyle.^[101] In planthoppers, jumps function both for escape and enhanced locomotion across vegetation, with leg synchronization aided by meshed gears in some species to prevent slippage.^[102] This catapult mechanism minimizes energy loss and maximizes acceleration, allowing insects as small as 5-10 mm to achieve accelerations up to 5,400 m/s² (approximately 550 times gravity). Jumping ability varies across Hemiptera, with reductions in groups adapted to non-terrestrial or specialized lifestyles. Aquatic forms, such as those in Corixidae, show diminished jumping capacity as propulsion shifts toward swimming, though some retain limited leaps for surfacing.^[103] Parasitic or highly sedentary taxa, including certain scale insects, lack robust hind leg modifications, relying instead on passive dispersal or minimal crawling during early instars.^[98] In pentatomid bugs like Erthesina fullo, terrestrial walking predominates with a tripod gait, but jumping is absent or rudimentary.^[104]

Aquatic adaptations and propulsion

Aquatic Hemiptera, particularly those in the suborder Heteroptera, have evolved specialized morphological and behavioral adaptations for effective locomotion in freshwater environments, enabling both submerged swimming and surface travel. In the Nepomorpha infraorder, species like water boatmen (Corixidae) and backswimmers (Notonectidae) rely on modified hind legs for propulsion, while semi-aquatic forms in Gerromorpha, such as velvet water bugs (Microveliidae), employ unique surface tension-based mechanisms. These adaptations allow them to navigate ponds, streams, and other habitats efficiently, balancing thrust generation with drag minimization at intermediate Reynolds numbers (typically 10–200).^[105] Submerged swimming in Corixidae involves synchronous rowing of the hind legs, which are transformed into oar-like paddles fringed with long setae. During the power stroke, the legs extend posteriorly at angles between 45° and 165°, with the tarsal setae splaying outward to maximize pressure drag on the water, generating the primary thrust—accounting for over 90% of forward propulsion. In the recovery stroke, the setae collapse against the leg, reducing drag as the limbs return anteriorly close to the body. This drag-based mechanism operates with a thrust efficiency of approximately 72%, though overall propulsive efficiency remains low at around 10% due to energy losses in unsteady flows.^[105]^[106] Backswimmers (Notonectidae) exhibit similar oar-like hind legs with dense fringes of hydrophobic hairs, but they swim inverted (belly-up) to position their pale ventral side downward for camouflage against the water surface. Propulsion combines leg rowing with undulating body motions, where the hind legs perform alternating or synchronous strokes to achieve rapid bursts; kinematic analyses reveal four primary patterns, including cruising and turning maneuvers modeled via Fourier series for precise leg trajectories. These insects attain burst swimming speeds of up to 0.5 m/s, facilitated by the hair fringes that enhance traction during strokes by increasing effective paddle area. Sexual dimorphism, with females typically larger than males, influences locomotion, as bigger bodies incur higher drag but support greater endurance in sustained swimming.^[107]^[108]^[109] Surface skating in smaller Gerromorpha species, such as Microvelia, utilizes Marangoni propulsion rather than direct leg paddling. These bugs secrete surfactants from salivary glands via the rostrum, creating a localized surface tension gradient (reducing tension from 72 to 49 dynes/cm behind the insect) that drives fluid flow and propels the body forward at peak speeds of 17 cm/s—roughly twice their walking speed on solid surfaces. This chemical gradient-based mechanism serves primarily as an escape response, allowing rapid directional changes without physical contact disrupting the water film.^[110] Diving adaptations in Nepomorpha enhance propulsion and survival during submerged excursions. Many species, including Notonectidae, trap air bubbles against the abdomen or under wings using hydrophobic hairs, forming a physical gill that extracts oxygen from surrounding water and extends dive times to over 2 minutes by replenishing 20% of consumed O₂ via diffusion. The hairy fringes on hind legs not only aid propulsion by providing traction against water resistance during strokes but also help retain air layers for buoyancy control, with haemoglobin in the hemolymph stabilizing bubble volume by releasing stored oxygen. These features enable efficient dives for foraging or evasion, minimizing the need for frequent resurfacing.^[111]^[111]

Ecology

Feeding strategies

Hemiptera exhibit diverse feeding strategies adapted to their piercing-sucking mouthparts, which enable them to extract liquid nutrients from plants, animals, or other insects. These strategies primarily fall into herbivory, predation, and hematophagy, each supported by specialized salivary secretions and gut modifications to process imbalanced or dilute diets.^[112] Herbivorous Hemiptera, particularly in the suborders Sternorrhyncha and Auchenorrhyncha, feed predominantly on phloem or xylem sap, which provides carbohydrates but is deficient in essential amino acids and other nutrients. Aphids (Aphididae), for example, insert their stylets into phloem sieve tubes to suck nutrient-rich sap, relying on endosymbiotic bacteria such as Buchnera aphidicola to synthesize essential amino acids from non-essential precursors in the diet. Similarly, xylem feeders like spittlebugs extract water-laden sap under negative pressure, compensating for its low nutrient density through high-volume intake and microbial symbionts.^[112]^[113]^[114]^[115] Predatory Hemiptera, mostly in the suborder Heteroptera, capture and immobilize prey using toxic saliva injected through their proboscis, which contains hydrolytic enzymes that paralyze victims and liquefy internal tissues for easier ingestion. Assassin bugs (Reduviidae) exemplify this strategy, secreting a cocktail of proteases, lipases, and neurotoxins to predigest arthropod prey externally before sucking up the resulting slurry. This extra-oral digestion enhances nutrient extraction efficiency, allowing predators to consume a wide range of invertebrates without solid food processing.^[116]^[117]^[118] Hematophagous species, also primarily Heteroptera such as bed bugs (Cimex lectularius) and kissing bugs (Triatominae), feed on vertebrate blood as ectoparasites, piercing skin to access capillaries and injecting anticoagulants in saliva to facilitate blood flow. These insects locate hosts using cues like carbon dioxide exhaled from breath, which activates antennal chemoreceptors and guides orientation toward potential meals. Blood provides a balanced nutrient profile, supporting reproduction and development, though feeding occurs infrequently due to the meal's high volume.^[119]^[120]^[121] To handle the excess water and sugars in sap or blood diets, many Hemiptera possess specialized gut structures for nutrient processing, with variations across suborders. Sap-feeders in Sternorrhyncha (e.g., whiteflies) and Auchenorrhyncha (e.g., leafhoppers) typically feature a filter chamber where the foregut and hindgut connect directly, rapidly shunting water and excess solutes past the midgut for excretion as honeydew while retaining nutrients. In contrast, aphids often lack a true filter chamber but achieve similar function through coiled hindgut segments adhering to the midgut, and predatory Heteroptera generally forgo such structures due to less watery diets. These adaptations prevent osmotic imbalance and optimize amino acid uptake.^[122]^[123]^[124]^[125]

Symbiotic relationships

Hemiptera exhibit diverse symbiotic relationships, particularly with bacteria and other insects, which play crucial roles in nutrition, reproduction, and protection. Endosymbiotic bacteria are prevalent in many hemipteran lineages, providing essential nutrients that supplement their specialized diets. For instance, in aphids (Aphididae), the obligate endosymbiont Buchnera aphidicola synthesizes essential amino acids, such as phenylalanine and tryptophan, which are scarce in the phloem sap that aphids feed on, enabling host survival and reproduction.^[126] Similarly, in some heteropterans, such as the lygaeoid bug Henestaris halophilus, the endosymbiont Candidatus Sodalis baculum supplies amino acids like tyrosine and lysine, along with cofactors, supporting the host's metabolic needs in nutrient-poor environments.^[127] Mutualistic interactions between Hemiptera and ants are widespread, often involving trophobiosis where hemipterans produce honeydew—a sugar-rich excretion from their feeding—that ants harvest in exchange for protection against predators. In aphid-ant associations, ants such as Lasius niger tend aphid colonies, removing waste and defending them from parasitoids and other insects, which can increase aphid population growth rates by up to 50% in tended colonies compared to untended ones.^[128] Scale insects (Coccidae and related families) form analogous mutualisms with ants, where species like Formica spp. protect scales from natural enemies while feeding on their honeydew, enhancing scale survival and reproduction in exchange for a reliable carbohydrate source.^[129] Certain endosymbionts in Hemiptera act as reproductive manipulators, with Wolbachia being a prominent example that induces cytoplasmic incompatibility in infected hosts. In various hemipteran species, including aphids like Acyrthosiphon pisum, Wolbachia infection causes embryonic lethality in crosses between infected males and uninfected females, but not in other combinations, thereby promoting the spread of infected lineages through maternal transmission.^[130] This manipulation favors Wolbachia-bearing females, altering host population dynamics and genetic diversity.^[131] The evolutionary history of hemipteran endosymbionts involves ancient acquisitions followed by extensive genome streamlining. Buchnera was acquired by the aphid lineage over 100 million years ago, with its genome reducing from an estimated 4-5 Mb ancestral size to around 0.6 Mb through gene loss, retaining primarily genes for essential amino acid biosynthesis while relying on host provisions for other functions.^[132] Similarly, Sodalis-like symbionts in heteropterans represent more recent acquisitions in some cases, but exhibit parallel genome reductions, losing non-essential genes while preserving metabolic pathways critical for host nutrition, reflecting co-evolutionary pressures in stable symbioses.^[133] Gut symbionts in certain Hemiptera further complement these relationships by aiding in digestion, though their roles are secondary to the primary nutritional endosymbionts.^[134]

Defensive adaptations and prey roles

Hemiptera employ a variety of chemical defenses to deter predators, particularly in the suborder Heteroptera, where adults possess metathoracic scent glands that produce volatile compounds such as aldehydes. These secretions, including trans-2-octenal and trans-2-decenal in species like the brown marmorated stink bug Halyomorpha halys, are released upon disturbance and serve as alarm pheromones and repellents, causing irritation to predators' sensory systems.^[135] In pyrrhocorids such as Dysdercus species, defensive secretions from dorsal abdominal glands in nymphs and metathoracic glands in adults include hydrocarbons and esters that contribute to aposematic signaling and predator aversion.^[136] Reflex bleeding, or autohaemorrhaging, occurs in some Hemiptera, where individuals exude hemolymph from intersegmental membranes or tarsi upon attack; in froghoppers (Cercopidae), this coagulates rapidly to entangle predator mouthparts, enhancing escape probability.^[137] Physical defenses in Hemiptera often involve mimicry and behavioral aggregation. Certain aphids, such as the root-dwelling Paracletus cimiciformis, exhibit aggressive mimicry where one morph chemically and morphologically imitates ants to infiltrate ant colonies, avoiding predation by exploiting ant social behaviors.^[138] Aggregation provides collective protection, as seen in aphid colonies like Aphis nerii, where clustered individuals amplify alarm pheromones and dilute per capita risk from predators, with studies showing reduced attack rates on grouped versus solitary aphids.^[139] In heteropterans like stink bugs, nymph aggregations facilitate synchronized defensive displays, including scent gland discharge, which overwhelms attackers.^[140] Camouflage and behavioral adaptations further aid predator avoidance in Hemiptera. Thanatosis, or feigning death, is observed in aphids, where disturbed individuals enter tonic immobility, reducing movement cues that trigger predator strikes; this response in species like Acyrthosiphon pisum can last minutes and correlates with higher survival against visual hunters.^[141] Escape jumps serve as rapid evasion tactics, particularly in planthoppers and froghoppers; for instance, the spotted lanternfly Lycorma delicatula uses unilateral hindleg structures to propel jumps up to several body lengths, powered by catapult-like mechanisms in the metathorax, allowing quick relocation to safer substrates.^[142] Froghoppers (Philaenus spumarius) similarly employ powerful jumps from plant surfaces by anchoring tarsi to prevent slippage, achieving accelerations over 300 m/s² for effective predator evasion.^[143] As prey, Hemiptera play integral roles in food webs, serving as a primary resource for predators like birds, spiders, and insects. Aphids, a dominant group, constitute a high-protein food source (up to 70% dry mass protein), supporting the diet of insectivorous birds, which collectively consume an estimated 400–500 million metric tons of arthropods annually, including substantial Hemiptera biomass.^[144]^[145] Spiders, particularly web-builders, frequently prey on aphids and other small hemipterans, with dietary analyses revealing Hemiptera as 10–30% of consumed biomass in agricultural habitats, linking hemipteran populations to higher trophic levels and influencing predator fitness through nutrient transfer.^[146] This prey role underscores Hemiptera's position as foundational herbivores in terrestrial and aquatic ecosystems, sustaining biodiversity via energy flow to vertebrates and invertebrates.^[147]

Relationships with humans

Agricultural and economic impacts

Hemiptera, particularly aphids and stink bugs, represent significant agricultural pests due to their feeding habits and ability to transmit plant pathogens. Aphids (Aphididae) are among the most economically important pests in temperate agriculture, causing direct damage through sap feeding and indirect harm by vectoring plant viruses such as soybean mosaic virus and alfalfa mosaic virus.^[148] Stink bugs, including species like the southern green stink bug (Nezara viridula) and the invasive brown marmorated stink bug (Halyomorpha halys), inflict damage by piercing seeds and fruits, leading to yield reductions and quality degradation in crops such as soybeans and tree fruits.^[149]^[150] For instance, N. viridula feeding on soybeans results in smaller seeds, discoloration, and wrinkling, with economic thresholds set at 36 bugs per 100 net sweeps to prevent significant losses.^[149]^[151] The economic impacts of these pests are substantial, with annual losses in the billions globally and notable figures in the United States. Aphid infestations, particularly soybean aphids, can reduce yields by up to 40% if unmanaged; for example, potential losses from aphid infestations such as the cereal grass aphid in the Pacific Northwest could reach about $120 million if damaging 5% of regional wheat production.^[152]^[153] A 2025 study estimated wheat disease losses, including those from aphid-vectored viruses, at $2.9 billion across the US and Canada from 2018 to 2021.^[154] The 2010 outbreak of the brown marmorated stink bug caused over $37 million in damages to mid-Atlantic apple producers, while N. viridula and related species drive consistent yield and quality losses in soybean production across the southern U.S.^[155]^[156] These invasive and native pests exacerbate costs through increased management expenses and reduced market value of affected commodities. In 2022, invertebrate pests including Hemiptera reduced soybean yields by 1.9% across 18 US states.^[157] Management of Hemiptera pests relies on integrated pest management (IPM) strategies that integrate biological, chemical, and cultural controls to minimize economic damage while reducing environmental risks. Biological controls include natural enemies such as predatory insects and parasitoids; for example, tachinid flies and big-eyed bugs (Geocoris spp.) target stink bugs, while lady beetles and lacewings suppress aphid populations.^[158]^[159] Chemical controls, like targeted insecticides, are used judiciously to avoid resistance and non-target effects, often applied based on economic thresholds such as 273 aphids per plant for soybeans.^[160] Cultural practices, including crop rotation, planting resistant varieties, and trap crops, help disrupt pest life cycles and reduce infestation levels in field crops.^[158]^[161] Certain Hemiptera species provide economic benefits through their role in IPM as predators of other pests. Minute pirate bugs (Orius spp.), such as Orius insidiosus, are key generalist predators that feed on aphids, thrips, and spider mites in greenhouses and field crops, often augmentatively released to enhance biological control efficacy.^[162]^[163] These beneficial bugs contribute to sustainable pest suppression, reducing reliance on chemical inputs and supporting higher yields in protected agriculture systems.^[164]

Medical and veterinary significance

Certain species of Hemiptera, particularly within the suborder Heteroptera, serve as significant vectors for diseases affecting humans, while others cause direct harm through bites or infestations. Kissing bugs, belonging to the Triatominae subfamily of the Reduviidae family (e.g., genera Triatoma and Rhodnius), are primary vectors of Trypanosoma cruzi, the protozoan parasite responsible for Chagas disease, a zoonotic infection that can lead to chronic cardiac and gastrointestinal complications if untreated.^[165]^[166] Transmission occurs when infected bugs defecate near the bite site during blood meals, allowing the parasite to enter through skin abrasions or mucous membranes; acute symptoms include fever and swelling, while chronic phases affect up to 30% of cases with severe organ damage.^[167] Bed bugs (Cimex lectularius and C. hemipterus, family Cimicidae) do not transmit diseases but pose medical risks through hematophagous feeding, triggering allergic reactions in sensitized individuals via salivary antigens.^[168] In veterinary contexts, Hemiptera impact livestock and poultry primarily through blood-feeding and associated stress or secondary infections. Kissing bugs transmit T. cruzi to wild and domestic mammals, including dogs and cattle, contributing to animal reservoirs that sustain human transmission cycles; canine Chagas disease manifests as cardiomyopathy, with seroprevalence reaching 10-20% in endemic US regions.^[166] Bed bugs infest poultry facilities, where heavy populations cause anemia, feather pecking, cloacal irritation, and reduced egg production in chickens; historical outbreaks in mid-20th-century farms led to significant economic losses before widespread insecticide use.^[169]^[170] Bites from these hemipterans commonly induce localized pruritus and erythematous papules due to histamine release, but severe responses include urticaria, bullous reactions, or anaphylaxis, particularly from repeated exposures to kissing bug saliva, which contains potent allergens like trypsin-like enzymes.^[171]^[172] In rare cases, massive bed bug infestations exacerbate asthma or cause iron-deficiency anemia in hosts.^[173] Control strategies emphasize integrated vector management, including surveillance through community reporting and timed manual collections to detect infestations, followed by targeted pyrethroid insecticide applications in domiciles and peridomiciles.^[174]^[175] Recent developments highlight Chagas disease's emergence as endemic in the United States, with locally acquired human cases reported in eight states since 2013 and increasing detections of infected Triatoma species in the 2020s, prompting enhanced CDC monitoring and public education.^[176]^[167]

Cultural and conservation aspects

Hemiptera have played notable roles in human culture through their derived products and as edible resources. The cochineal scale insect, Dactylopius coccus, serves as the source of carmine, a vibrant red dye extracted from the dried bodies of females, historically prized by Mesoamerican civilizations and later exported to Europe following the Spanish conquest of Mexico in the 16th century.^[177] This dye, yielding up to 20% of the insect's dry weight as pigment, has been used in textiles, cosmetics, and food coloring, with production still centered in Peru and the Canary Islands.^[178] Similarly, lac bugs of the genus Kerria, particularly K. lacca, secrete shellac, a natural resin harvested from infested host trees in India and Thailand, processed into a versatile material for varnishes, polishes, and pharmaceuticals due to its adhesive and insulating properties.^[179] Certain Hemiptera species are consumed as food in various cultures, valued for their nutritional content. In East and South Asia, cicadas such as Cryptotympana atrata are harvested and prepared by frying or roasting, providing high protein levels (around 21.4 g per 100 g dry weight) and low fat, alongside essential amino acids and minerals that contribute to dietary diversity.^[180] In Mexico, the eggs of water boatmen (Corixidae family), known as ahuautle or "Mexican caviar," have been a pre-Hispanic delicacy since Aztec times, harvested from lakes and noted for their rich macronutrient profile, including proteins (up to 58% dry weight), lipids, and phenolic compounds with antioxidant activity.^[181] Hemiptera feature prominently in cultural symbolism, particularly cicadas. In Chinese art and poetry, cicadas symbolize rebirth, purity, and integrity, appearing in Tang dynasty verses as emblems of moral steadfastness and in jade carvings placed in tombs to invoke immortality and transformation.^[182] In Greek folklore, cicadas were associated with music and the divine, linked to myths like that of Tithonus, whose eternal life without youth transformed him into a cicada, representing the insect's song as a poetic echo of human transience.^[183] Conservation efforts for Hemiptera are challenged by multiple threats, including habitat loss from urbanization and agriculture, which fragments ecosystems essential for species like scale insects and cicadas.^[184] Pesticide use exacerbates declines by reducing populations of non-target species, while climate change alters distributions, potentially contracting ranges for temperate Hemiptera and expanding invasive pests like stink bugs northward.^[185] Globally, invasive species including Hemiptera have cost agriculture over $644 billion from 1970 to 2020.^[186] Although no Hemiptera are currently listed as endangered by the IUCN, several periodical cicada broods (Magicicada spp.) are classified as Near Threatened due to habitat pressures, underscoring the need for protected areas to safeguard biodiversity.^[187]

References

[1]
Orders of Insects: Hemiptera – Insect Science - Open Textbooks @ UQ
Three Suborders: · Includes whiteflies, aphids, coccids, mealybugs, scale insects; Mouthparts appear to arise from between first pair of legs (Sternorrhyncha ...Order Hemiptera: Bugs... · Three Suborders · 30 Orders Of Insects...Missing: taxonomy | Show results with:taxonomy
[2]
Order Hemiptera Suborder Heteroptera - ENT 425
Members of the suborder Heteroptera are known as “true bugs”. They have very distinctive front wings, called hemelytra, in which the basal half is leathery and ...
[3]
True bugs (Order: Hemiptera) - Amateur Entomologists' Society (AES)
Information on the True bugs (Order: Hemiptera). Including shieldbugs, plant bugs, bed bugs, pondskaters, cicadas, water bugs, aphids and scale insects.Missing: ecology | Show results with:ecology
[4]
[PDF] Bio Ecology of Aquatic and Semi-Aquatic Insects of Order Hemiptera ...
Nov 23, 2021 · Background: Hemiptera is an order of insects, commonly called true bugs, comprising over 80,000 species. Most hemipterans feed on plants.
[5]
Phylogeny and evolution of hemipteran insects based on expanded ...
Sep 2, 2024 · Traditionally, Hemiptera has been classified into four major suborders: Sternorrhyncha (aphids, scale bugs, whiteflies, and psyllids), ...
[6]
Phylogenetics and Higher Classification - WordPress at UD |
The suborders of Hemiptera remain Sternorrhyncha, Auchenorrhyncha, Heteroptera and Coleorrhyncha (or Prosorrhyncha for the combined grouping), despite ongoing ...
[7]
Aphididae | INFORMATION | Animal Diversity Web
Within the family Aphididae, also known as aphids or plant lice, there are 24 subfamilies, 510 genera, and 5109 species. Aphids are very small insects that live ...
[8]
Cicada | Sound, Shell, Life Cycle, Map, & Facts | Britannica
Oct 14, 2025 · Cicadas form a large family of insects, with over 3,000 species. Cicadas are found in tropical and temperate areas worldwide, including deserts, ...
[9]
Number of Genera & Species - North Dakota State University
The total number of genera in Pentatomidae is 896, and the total number of species is 4722.
[10]
Assassin bug | Description, Scientific Name, Reduviidae, Insect ...
Sep 26, 2025 · An assassin bug is any of about 7000 species of true bugs that are characterized by a thin necklike structure connecting the narrow head to ...
[11]
A revised classification of the assassin bugs (Hemiptera: Heteroptera
Jul 17, 2024 · Our new classification for Reduviidae reduces the number of subfamilies to 19 and recognizes 40 tribes. We describe three new subfamilies to ...Missing: suborders | Show results with:suborders
[12]
https://www.mapress.com/zt/article/view/40419
[13]
A report on newly described taxa of Hemiptera and other 28 orders ...
Results: From 2022 to 2023, a total of 727 new taxa were published for Hemiptera and 28 other insect orders in China, comprising 668 new species/subspecies, 58 ...
[14]
Phylogenomics and the evolution of hemipteroid insects - PNAS
Nov 26, 2018 · We also recovered monophyly of most currently recognized suborders, infraorders, and superfamilies within these groups as well as resolving ...
[15]
Phylogeny and evolution of hemipteran insects based on expanded ...
Sep 2, 2024 · We constructed multiple datasets comprising 298 Hemiptera species, covering all four suborders, 13 infraorders, 27 superfamilies, and 85 ...
[16]
Diversification and extinction of Hemiptera in deep time - Nature
Mar 3, 2025 · This study leverages birth-death models in a Bayesian framework and the fossil record of Hemiptera to estimate their past diversity dynamics.
[17]
Bayesian modelling of the fossil record enlightens the evolutionary ...
Sep 10, 2025 · Hemiptera, the fifth most diverse insect order, are characterized by their high diversity in deep time, with 145 known extinct families.
[18]
Fossils reshape the Sternorrhyncha evolutionary tree (Insecta ...
Jul 9, 2020 · The order Hemiptera is subdivided into six suborders—extinct Paleorrhyncha (archescytinoids), Sternorrhyncha (modern aphids, scale insects ...
[19]
New hemipteran insects (Hylicellidae, Ipsviciidae, Heteroptera) from ...
Jun 2, 2025 · Four new hemipteran insects are described from the Triassic (Norian) Cow Branch Formation of Virginia and North Carolina (USA)
[20]
(PDF) The first fossil Dipsocoridae found in the early Eocene amber ...
Aug 6, 2025 · The first fossil Dipsocoridae found in the early Eocene amber of France (Hemiptera: Heteroptera) · Abstract and Figures · Citations (7) ...
[21]
Why so many Hemiptera invasions? - Liebhold - Wiley Online Library
Sep 6, 2024 · About 100,000 hemipteran species are described worldwide, placing this order fifth in global species richness among insect orders after ...
[22]
[PDF] How Many Species of Insects and Other Terrestrial Arthropods Are ...
Sep 23, 2017 · ... species of Hemiptera, the world total would be 184,000–193,000. They suggested that Hemiptera constitute 7.5–10% of the canopy fauna, and ...
[23]
Ultraconserved elements from transcriptome and genome data ...
May 29, 2024 · Sternorrhyncha, one of the four major suborders of Hemiptera, is a phytophagous taxon inclusive of nearly 18 000 described species.
[24]
Suborder Auchenorrhyncha - NC State University Libraries
Dec 1, 2008 · The suborder may be divided into two major groups (treated as infraorders by some workers) and four superfamilies as follows: infraorder ...<|control11|><|separator|>
[25]
Heteroptera – IHS - Internationa Heteropterists' Society
Jan 25, 2022 · Heteroptera, or true bugs, are a diverse suborder of Hemiptera with over 45,000 species. They have flat wings, a piercing-sucking mouth, and ...
[26]
Hemiptera collection | MNHN - Muséum national d'Histoire naturelle
Depending on the taxonomic group, 30 to 70% of described 100,000 Hemiptera species are represented there, making it an unrivalled source of primary information.
[27]
Spatial Patterns and Determinants of the Diversity of Hemipteran ...
Large-scale patterns of species richness is an important issue in biogeography and ecology. The Qinghai-Tibetan Plateau (QTP) is a biodiversity hotspot in ...
[28]
[PDF] 中国半翅目等29目昆虫新分类单元报告(2022‒2023) - 生物多样性
... new taxa were published for Hemiptera and 28 other insect orders in China, ... Specifically, Hemiptera had 252 new species,. 22 new genera, and 1 new ...
[29]
Aphids Management Guidelines--UC IPM
### Summary of Myzus persicae (Green Peach Aphid), Aphididae Traits
[30]
[PDF] Aphids & Their Relatives - New Mexico Department of Agriculture
Scientifically: The insects in this fact sheet are all members of the Order. Hemiptera, Suborder Sternorrhyncha. ... Close-up of San Jose scale, Quadraspidiotus.
[31]
The Adipokinetic Peptides of Hemiptera: Structure, Function, and ...
There are more than 12 000 known species of planthoppers, that feed off plant sap as adults and nymphs, thereby producing honeydew which is often related with ...
[32]
Brown Marmorated Stink Bug
### Summary of Brown Marmorated Stink Bug (Pentatomidae)
[33]
Chapter 3: Entomology - Pressbooks at Virginia Tech
Insect Form and Structure—Morphology. All adult members of class Insecta are identifiable by having three body regions, three pairs of legs, and one pair of ...
[34]
Aphididae - an overview | ScienceDirect Topics
The family Aphididae comprises more than 4300 species, all of which are specialized to feed on phloem sap [1]. Aphids can have a dramatic negative impact on ...
[35]
True Bugs - Ask A Biologist - Arizona State University
Oct 6, 2010 · They can range in size from very small (~1mm), such as bed bugs, to huge (15cm/6 inches), like the giant water bug.
[36]
Hemiptera - bugs, aphids, cicadas - ento.csiro.au
The true bugs have forewings that are hardened at the base and membranous at the tips. They sit flat over the abdomen hiding the membranous hind wings. The head ...Missing: morphology | Show results with:morphology
[37]
Wing dimorphism in aphids | Heredity - Nature
Jul 5, 2006 · In most species the sexual females are wingless, with exceptions in more primitive taxa. Sexual males of Eriosomatinae, Anoeciinae, ...
[38]
Hemiptera: Morphology and Phylogeny - Longdom Publishing
The Hemiptera are called 'true' bugs comprising over 80,000 species within groups such as the cicadas, aphids, plant hoppers, leafhoppers, bed bugs, and shield ...Missing: plan | Show results with:plan
[39]
Chapter 4 Classes of Arthropod Pests of the Urban Community
Aug 23, 2002 · A pair of scent glands on each segment, opening laterally, discharge a repugnant secretion. In some species, it can be discharged to a ...
[40]
Morphology of Insects <Biological Control
The head capsule has become a highly evolved or specialized structure involving at least five primitive or generalized metameres. The first metamere or ...<|control11|><|separator|>
[41]
The structure of extremely long mouthparts in the aphid genus ...
One of the most striking characters associated with the Hemiptera is a set of modifications in their mouthparts (labium, labrum, maxillae and mandibles), which ...Missing: origin | Show results with:origin
[42]
Unique Fine Morphology of Mouthparts in Haematoloecha nigrorufa ...
Jun 22, 2020 · The mouthpart complex in Hemiptera is often called the “rostrum”, “sucking beak”, or piercing-sucking mouthparts [10].
[43]
Salivary Proteins of Russian Wheat Aphid (Hemiptera: Aphididae)
Feb 1, 2010 · Soluble saliva is thought to be composed of pectinases, cellulases ... Our study is the first to detect this enzyme in aphid saliva.
[44]
Study of the Salivary Glands in Triatominae (Hemiptera, Reduviidae ...
The salivary glands of triatomines are important to hematophagy because their saliva is rich in anticoagulant and hemolytic proteins. The salivary glands of ...
[45]
Mitochondrial phylogenomics of Hemiptera reveals adaptive ...
Sep 6, 2017 · More recently, Hemiptera has been subdivided into four major suborders, Sternorrhyncha (Psylloidea, Aleyrodoidea, Aphidoidea, and Coccoidea) (e ...
[46]
Phylogenomics and the evolution of hemipteroid insects - PMC - NIH
Nov 26, 2018 · The origin of this group apparently coincided with a distinct shift in mouthpart morphology and feeding habits toward piercing and sucking.
[47]
[PDF] Getting to know the insects
True bugs (Hemiptera = half winged) – forewings, called hemelytra (the basal half is hardened and the distal half is membranous), and membranous hindwings are ...
[48]
[PDF] Hemiptera, Heteroptera - Insecta - eScholarship
The fore-wing (hemelytra or hemielytra) is usually hard and leathery at the basal half and membranous at the distal part.
[49]
Hemiptera – Heteroptera – UNBC BIOL 322, Entomology
The suborder Heteroptera, the true bugs, are so-named because their forewings are partially sclerotized at the base, forming hemelytra. In addition to this ...
[50]
(PDF) Hemíptera: Auchenorrhyncha - ResearchGate
Apr 13, 2024 · ... tegmina (in most taxa); and wings with pairs of. anal veins forming a set of Y-shaped veins on the distal margin. (in most taxa) (BAPTISTA ...
[51]
From micropterism to hyperpterism: recognition strategy and ... - NIH
A standardized and homology-driven groundplan terminology for tegmina types, structures and vein patterns in Hemiptera Fulgoromorpha, including fossils, is ...
[52]
Identifying Differential Gene Expression in Wing Polymorphism ... - NIH
Dec 10, 2018 · This is particularly observed in terms of wing length, including macropterous, brachypterous, and apterous morphs, owing to differences in the ...
[53]
Molecular Mechanisms of Wing Polymorphism in Insects
Oct 12, 2018 · We first introduce the wing polymorphisms of planthoppers, aphids, and crickets and describe the environments that result in the production of ...
[54]
Aphid polyphenisms: trans-generational developmental regulation ...
A number of environmental cues affecting the dispersal phenotypes (winged/macropterous or wingless/brachypterous) of aphids have been identified, especially in ...
[55]
Aphid wing dimorphisms: linking environmental and genetic control ...
Here, I review investigations into molecular mechanisms underlying phenotypic plasticity in the aphid wing dimorphism system.
[56]
Differential wing polyphenism adaptation across life stages under ...
Jun 19, 2019 · Wing polyphenism is essential for the aphid life cycle and, by allowing for migration to fresh food resources, it may contribute to determining ...
[57]
Preliminary Study of Wing Interference Patterns (WIPs) in Some ...
Jul 30, 2013 · The fore wings of scale insect males possess reduced venation compared with other insects and the homologies of remaining veins are ...
[58]
Nomenclature of the veins of the fore wings of male scale insects ...
Dec 19, 2022 · Compared with psyllids and aphids, the remaining wing venation in male scale insects is much more reduced, usually with two well-developed veins ...
[59]
(PDF) The systematics of the Hemiptera - ResearchGate
In this article the main phylogenetic hypotheses for Hemiptera and its subgroups are reviewed. Important references regarding catalogs and identification aids ...
[60]
Wing base structure supports Coleorrhyncha + Auchenorrhyncha ...
Aug 7, 2025 · Wing base structure supports Coleorrhyncha + Auchenorrhyncha (Insecta: Hemiptera) ... wing folding is a key for the shelter function of the elytra ...
[61]
Guide To Insect Orders and Families <Legner, E. F.
Antennae short, bristle-like, and inconspicuous; abdomen with 2-3 thread-like caudal filaments; wings with numerous veins and cells; halteres absent; usually >5 ...
[62]
Diptera – Introduction – UNBC BIOL 322, Entomology
The hind wings of Diptera are reduced and modified into small, club-like structures called halteres. Halteres are used as a balance organ to facilitate rapid, ...
[63]
Form-function relationships of the compound eyes and sensory ...
Form-function relationships of the compound eyes and sensory sensilla of a tiny arboreal hemipteran herbivore: Adaptations for close encounters with leaves.Abstract · 3.1. Compound Eye · 3.3. 5. Sensilla Basiconica
[64]
Evidence for trichromacy in the green peach aphid, Myzus persicae ...
Under the chosen adaptation conditions, the green peach aphid showed three peaks of spectral sensitivity, one in the green region around 530 nm, a secondary ...
[65]
Visual Detection and Avoidance of Pathogenic Bacteria by Aphids
Oct 8, 2018 · Pea aphids use vision in the presence of ultraviolet light to avoid pathogenic bacteria. Avoidance is mediated by visual detection of fluorescence from ...
[66]
Morphological characterization of the antennal sensilla of the Asian ...
The filiform antennae of both sexes were of the conventional type comprised ... S.E. Halbert et al. Asian citrus psyllids (Sternorrhyncha: Psyllidae) ...
[67]
Environmental Correlates of Sexual Signaling in the Heteroptera
Insect antennae often host pheromone signal receptors and have been found to coevolve with the olfactory system, developing unique and specific receptors ...
[68]
Communication by substrate-borne mechanical waves in insects
Biotremology is the study of substrate-borne vibrations used by insects for communication, detected by specialized organs and used for inter- and intraspecific ...
[69]
Coordinative mediation of the response to alarm pheromones by ...
Aphids release alarm pheromones, mainly (E)-β-farnesene (EβF), to alert nearby aphids when they are attacked by predators or parasitoids (Nault et al., 1973; ...
[70]
Male orientation to trail sex pheromones in parasitoid wasps
brassicae, females are gathered at emergence sites, their trail sex pheromone ... In this species, males and females often emerge from different aphid ...
[71]
The Role of the Tymbal in Cicada Sound Production
Apr 1, 1995 · These sounds are generated by a pair of tymbals, which are relatively small stiff membranes, located dorsolaterally on the anterior part of the abdomen.ABSTRACT · INTRODUCTION · MATERIALS AND METHODS · RESULTS
[72]
COMMUNICATION WITH SUBSTRATE-BORNE SIGNALS IN ...
Oct 8, 2002 · Stridulation, the process whereby sound or vibration is produced by friction of two body parts moving across one another, is widespread among ...
[73]
[PDF] Acoustical Communication in Heteroptera (Hemiptera
Sep 14, 2011 · Vibrational signals and songs are produced by stridulation (stridulatory device, stridulatory organs), by body vibration, or by a simple tymbal ...<|control11|><|separator|>
[74]
The role of the tymbal in cicada sound production - PubMed
Contraction of the tymbal muscle causes the tymbal plate to swing inwards, acting as a lever so that the surface of the tymbal moves through more than twice the ...
[75]
Evolution of stridulatory mechanisms: vibroacoustic communication ...
Apr 26, 2023 · Stridulation occurs when two sclerotized structures rub together to produce vibratory and acoustic (vibroacoustic) signals, such as a cricket ' ...
[76]
Sound-producing mechanisms of Buenoa macrotibialis Hungerford ...
The stridulatory mechanism composed of the tibial combs and rostral prongs is involved in producing trills, chirps, and dart-chirps.
[77]
Vibrational behavior of psyllids (Hemiptera: Psylloidea)
In conclusion, the signal producing mechanism of psyllids may involve two sets of morphological structures. The first is stridulation between the axillary ...
[78]
Wing buzzing as a mechanism for generating vibrational signals in ...
Jan 24, 2024 · We argue that males of C. pyrisuga produce the vibrations primarily by wing buzzing, that is, tremulation that does not involve friction between the wings and ...
[79]
The evolution of insect metamorphosis: a developmental and ... - PMC
Aug 26, 2019 · We believe that the hemimetabolous forms also have a three-part life history of pronymph, nymph and adult, although one phase is cryptic within ...1. Introduction · (a). Developmental... · 5. Genetic Circuits...Missing: diapause | Show results with:diapause
[80]
Aphids, Cicadas, Scales, Leafhoppers, and others
Adult cicadas damage twigs and the subterranean nymphs feed on tree roots. Some cicada species have a long life-cycle, with the nymphs taking 17 years to grow.Missing: examples diapause
[81]
Aphids, in-depth - Wisconsin Horticulture
Most aphids are about 1/8 inch long and all are soft-bodied. Their pear-shaped bodies have “exhaust pipes” (cornicles) protruding from the back end of the ...
[82]
Effects of Diapause on Halyomorpha halys (Hemiptera - PubMed
Aug 11, 2018 · Diapause and cold tolerance can profoundly affect the distribution and activity of temperate insects. Halyomorpha halys (Stål) (Hemiptera ...
[83]
Acoustic signals in cicada courtship behaviour (order Hemiptera ...
Feb 28, 2006 · During pair formation, cicadas produce acoustic signals that allow sexual partners to meet. The male is generally the emitter, ...
[84]
Traumatic insemination and sexual conflict in the bed bug Cimex ...
We conclude that traumatic insemination is probably a coercive male copulatory strategy that results in a sexual conflict of interests.
[85]
[PDF] Morphology, Ontogeny, Reproduction, and Feeding of True Bugs
Most heteropterous males apparently deposit sperm in the female in the shape of a spermatophore; the sperm is then stored by the female in the spermatheca.<|control11|><|separator|>
[86]
(PDF) Egg attendance and brooding by males of the giant water bug ...
Aug 7, 2025 · Lethocerus mediusmales apparently remain with their mates through oviposition and are consistently found in attendance of eggs after the female ...
[87]
Aphid polyphenisms: trans-generational developmental regulation ...
In other words, cyclical parthenogenesis in aphids has evolved in order to facilitate asexual viviparous reproduction as well as by overwintering by diapausing ...Missing: mating | Show results with:mating
[88]
Egg retention, viviparity and ovoviviparity in Paraneoptera
Aug 3, 2021 · The most studied viviparous species of Heteroptera is the polyctenid Hesperoctenes fumarius (Westwood, 1874), an ectoparasite of bats. The ...
[89]
THE ORGANIZATION OF FLIGHT MUSCLE IN AN APHID ...
THE ORGANIZATION OF FLIGHT MUSCLE IN AN APHID, MEGOURA VICIAE (HOMOPTERA). With a Discussion on the Structure of Synchronous and Asynchronous Striated Muscle ...
[90]
Asynchronous Muscle: A Primer - Company of Biologists journals
Sep 15, 2000 · Asynchronous muscles are high-frequency muscles. The wing-stroke frequency during flight of insects using asynchronous muscle ranges from ...Introduction · Features of the beetle muscle · Delayed stretch activation...
[91]
Differential expression of genes in the alate and apterous morphs of ...
Aug 31, 2016 · In aphids, alate and apterous morphs differ in presence or absence of wings and also differ in body structure, fecundity, longevity and behavior ...
[92]
Long-range aerial dispersal of cereal aphids as virus vectors in ...
This paper reviews studies on the reinvasion phenomenon including the recent 'Pests and Weather' project in Illinois (1983-1985) in which radar and traps ...
[93]
A TRADE‐OFF BETWEEN FLIGHT CAPABILITY AND ...
Mar 1, 2000 · Here we examine the potential cost of flight capability in males of the salt-marsh-inhabiting planthopper Prokelisia dolus (Hemiptera: ...
[94]
Reproductive cost of flight capability: a comparison of life history ...
1. Reproductive costs associated with flight capability were evaluated in the wing dimorphic planthopper, Prokelisia dolus Wilson, by comparing the life history ...
[95]
Probing Behavior of Apterous and Alate Morphs of two Potato ...
Dec 2, 2011 · Secondary host plant colonization by aphids involves alate and apterous morphs to spread in the population at a large scale by flying or, ...
[96]
"Nocturnal Flights of TRIATOMA (Hemiptera: Reduviidae) in Sabino ...
Triatoma flights appear to be due to starvation. If adult bugs find themselves without a wood rat host, they become starved within approximately 15 days ...
[97]
Hemiptera (bugs)
There are four main types of leg modifications that reflect bug habits; 1) cursorial for running where the femora and tibiae are elongate and slender; 2) ...
[98]
ADAPTATIONS IN SCALE INSECTS - Annual Reviews
In sedentary armored scale insects (Diaspididae), the antennae are entirely reduced to stumpy tubercles (10), whereas free-moving coccoids have long antennae ...Missing: locomotion | Show results with:locomotion
[99]
Jumping performance of froghopper insects
Dec 1, 2006 · In a jump, the attitude of the body is set by the front and middle legs, and the propulsion is delivered by rapid and synchronous movements of ...
[100]
Froghoppers jump from smooth plant surfaces by piercing them with ...
Froghoppers produce traction by piercing plant surfaces with sharp metal-enriched spines on their hind legs, deforming the cuticle plastically and leaving ...
[101]
Jumping strategies and performance in shore bugs (Hemiptera ...
Jan 1, 2009 · Jumping is a common means of escaping rapidly from predators or increasing the speed of locomotion. Many insects that jump can also fly and the ...Missing: responses | Show results with:responses
[102]
[PDF] Jumping performance of planthoppers (Hemiptera, Issidae)
jumps by flicking its abdomen forwards and then backwards while extending the tibiae and depressing the femora of the hind and middle legs. (Burrows and Morris, ...
[103]
Corixidae - an overview | ScienceDirect Topics
“Aquatic Hemiptera” is therefore synonymous with “aquatic Heteroptera. ... ability to jump and conceal themselves in crevices frustrates the best of collectors.
[104]
Interleg coordination in free-walking bug Erthesina fullo (Hemiptera
The pentatomid bug Erthesina fullo exhibits a tripod gait, when walking freely on horizontal substrates, like many other insects. In this study, amputations ...
[105]
The hydrodynamics of swimming at intermediate Reynolds numbers ...
Aug 1, 2014 · The present study aimed to resolve which forces create thrust and drag in the paddling of the water boatman (Corixidae), an animal that spans ...
[106]
Hydrodynamics of swimming in the water boatman, Cenocorixa bifida
Aug 6, 2025 · The animal is propelled by the synchronous rowing action of its hind legs. The propulsive cycle consists of a power stroke and a recovery stroke ...
[107]
Motion Patterns and Kinematic Modeling of Backswimmer Hind Legs
Nov 4, 2025 · This paper used three-dimensional high-speed cameras to extract the key points on the hind legs. The hind leg motion laws and the deformation ...Missing: per | Show results with:per
[108]
(PDF) Swimming Ability of Notonectidae (Hemiptera) - ResearchGate
Aug 10, 2025 · SWIMMING ABILITY OF NOTONECTIDAE ; (HEMIPTERA) ; By Steven H. · Gittelman* ; Biological Sciences Group, University of Connecticut ; Storrs, Conn.
[109]
Sex on the Beach: Sexual Size Dimorphism Among Aquatic Bugs ...
Jun 5, 2025 · Morphological differences between the sexes are generally minor, except that female body size is usually longer than that of males.
[110]
[PDF] WALKING ON WATER: Biolocomotion at the Interface
Peak speeds during Marangoni propulsion for Microvelia are approximately 17 cm/s, or twice the peak walking speed (Andersen 1982). By dusting the water ...
[111]
Gas exchange and dive characteristics of the free-swimming ...
Nov 1, 2015 · Many aquatic insects utilise air bubbles on the surface of their bodies to supply O2 while they dive. The bubbles can simply store O2, as in ...<|separator|>
[112]
Heritable symbiosis: The advantages and perils of an evolutionary ...
Aphids and other sap-feeding insects rely on phloem sap or xylem sap as their only food, and these diets are extremely limited in essential amino acids and ...Missing: strategies | Show results with:strategies
[113]
[PDF] development of genomic resources to study aphid response
Plant phloem sap contains high levels of sucrose and an unbalanced mixture of amino acids, and aphids rely on their endosymbiotic bacteria, Buchnera aphidicola, ...
[114]
Bacteriocyte Reprogramming to Cope With Nutritional Stress in a ...
Oct 25, 2018 · All phloem-feeding hemipterans are capable of utilizing phloem sap thanks to their intimate association with symbiotic bacteria which furnish ...
[115]
[PDF] David W. Tonkyn and Robert F. Whitcomb
Two large orders of insects-the Hemiptera and Homoptera-possess mouthparts that are highly modified for piercing tissues and extracting their fluid contents.
[116]
An Assassin's Secret: Multifunctional Cytotoxic Compounds in the ...
Apr 20, 2023 · Predatory assassin bugs produce venomous saliva that enables them to overwhelm, kill, and pre-digest large prey animals.
[117]
Paralyzing and Liquefying Venom of The Assassin Bug ... - NIH
Assassin bug venom has features in common with venoms from other animals, such as paralyzing and lethal activity when injected, and a molecular composition ...
[118]
EXTRA-ORAL DIGESTION IN PREDACEOUS TERRESTRIAL ...
The insects alternately pump the potent hydrolytic saliva and suck the liquified prey parts, delivering the nutrients to the digestive tract (22, 23).
[119]
Bedbugs and Infectious Diseases - PMC - PubMed Central - NIH
Bedbugs are brown and flat hematophagous insects. The 2 cosmopolite species, Cimex lectularius and Cimex hemipterus, feed on humans and/or domestic animals, and ...
[120]
The Olfactory Chemosensation of Hematophagous Hemipteran Insects
Aug 9, 2021 · Kissing bug nymphs are attracted by CO2-free traps baited with three host-odor components (ammonia, L-(+)-lactic acid, and hexanoic acid) ...
[121]
The Role of Antennae in Heat Detection and Feeding Behavior ... - NIH
Oct 31, 2020 · As in other hematophagous insects, bed bugs are capable of detecting and responding to CO2, odors, and heat. Carbon dioxide has previously been ...
[122]
Three-dimensional reconstruction of a whole insect reveals its ...
We report on the internal 3D structures of the brown planthopper, Nilaparvata lugens (Hemiptera: Delphacidae) at nanometer resolution for the first time.Missing: variations | Show results with:variations<|control11|><|separator|>
[123]
Whitefly–Plant Interactions: An Integrated Molecular Perspective
Oct 10, 2023 · The whitefly filter chamber shunts excess water and some solutes from the an- terior digestive system directly into the hindgut, while the ...
[124]
The Paraneopteran Orders - ENT 425
Excess water, sugars, and certain amino acids bypass most of the midgut and are shunted directly into the hindgut for excretion as honeydew.
[125]
[PDF] Identification and characterization of functional aquaporin water ...
Some hemipteran xylem and phloem-feeding insects have evolved specialized alimentary structures or filter chambers that rapidly transport water for excretion ...
[126]
Trading amino acids at the aphid–Buchnera symbiotic interface
Jul 23, 2019 · A transporter capable of transporting nonessential amino acids across the symbiosomal membrane of the pea aphid Acyrthosiphon pisum.
[127]
A New Bacteriome-Associated Endosymbiont of the Lygaeoid Bug ...
Oct 1, 2017 · Ca. Sodalis baculum is a mutualistic endosymbiont capable of supplying the amino acids tyrosine, lysine, and some cofactors to its host.
[128]
Ecological consequences of interactions between ants and ...
Oct 31, 2006 · The mutualistic interactions between ants and honeydew-producing hemipterans may dramatically increase the effects of ants as predators, ...
[129]
what benefits for Melissotarsus emeryi ants and armored scale ...
Jul 25, 2017 · Mutualisms between ants and sap-sucking insects generally involve clear benefits for both partners: the ants provide protection in exchange ...
[130]
Molecular characterization and genetic diversity of Wolbachia ...
Sep 28, 2023 · Wolbachia, a Gram-negative alpha-proteobacterium, is the most prevalent endosymbiotic bacterial group, associated with over 60% of insect ...<|control11|><|separator|>
[131]
A rapidly spreading deleterious aphid endosymbiont that uses ...
Apr 24, 2023 · Transfers of Wolbachia endosymbionts have been adopted to control arbovirus transmission by mosquitoes (23–25).
[132]
Genome Evolution of the Obligate Endosymbiont Buchnera aphidicola
Apr 15, 2019 · Collectively, these results indicate that extensive genome reduction occurred in the ancestral Buchnera prior to aphid diversification and that ...
[133]
Swapping symbionts in spittlebugs: evolutionary replacement of a ...
Jan 9, 2014 · Strikingly, the newly acquired Sodalis-like symbiont retains genes underlying efficient routes of energy production, including a complete TCA ...
[134]
Evolutionary loss and replacement of Buchnera, the obligate ... - NIH
Jan 23, 2018 · One evolutionary solution is the acquisition of a novel symbiont with a functionally intact genome. Whereas almost all aphids host the anciently ...
[135]
Alarm Odor Compounds of the Brown Marmorated Stink Bug Exhibit ...
Trans-2-octenal and trans-2-decenal are the major alarm aldehydes responsible for the scent of Halyomorpha halys, the brown marmorated stink bug.
[136]
Defensive secretions of nymphs and adults of five species of ...
The chemical composition of the defensive secretions of the nymphs and the adults of Dysdercus fasciatus was investigated. Thirty compounds were identified ...Missing: Hemiptera | Show results with:Hemiptera
[137]
Reflex Bleeding in Froghoppers (Homoptera: Cercopidae)
Reflex bleeding in New World froghoppers is described in detail for the first time. Prosapia sp. n. adults exude hemolymph from rupture lines in pretarsal pads ...Missing: pyrrhocoridae | Show results with:pyrrhocoridae
[138]
Aggressive mimicry coexists with mutualism in an aphid - PMC
Our results show that chemical mimicry is used by flat morph aphids to deceive worker ants and infiltrate their brood chamber. First, results of adoption tests ...
[139]
Collective Defense of Aphis nerii and Uroleucon hypochoeridis ...
Apr 29, 2010 · The prevalent way aphids accomplish colony defense against natural enemies is a mutualistic relationship with ants or the occurrence of a ...
[140]
Aggregation Behavior of the Southern Chinch Bug (Hemiptera
Aug 1, 2012 · The gregarious feeding behavior of the immature stage may aid in overcoming plant defenses and/or decreasing predation risk. An attractive ...
[141]
Aphids Playing Possum – Defensive or Mutualistic Response? - NIH
Jan 18, 2018 · Thanatosis, or the so called “death feigning” or “playing possum” is a well-recognized phenomenon in many animal taxa (Honma et al. 2006). It is ...
[142]
The Unilateral Jumping Structures of the Spotted Lanternfly ...
Jul 6, 2025 · Among arthropods, many Hemiptera insects exhibit exceptional jumping abilities. The jump of these insects is executed by metathoracic muscles, ...
[143]
Froghoppers jump from smooth plant surfaces by piercing them with ...
Feb 4, 2019 · Some of the fastest and most powerful jumps are performed by plant sap-sucking bugs of the order Hemiptera, which includes froghoppers, ...
[144]
Insectivorous birds consume an estimated 400–500 million tons of ...
Jul 9, 2018 · We estimate the prey biomass consumed by the world's insectivorous birds to be somewhere between 400 and 500 million metric tons year −1.Missing: nutritional | Show results with:nutritional
[145]
Arthropod prey vary among orders in their nutrient and exoskeleton ...
Dec 14, 2021 · However, the considerable variation in nutrient content observed among arthropod taxa (10%–85% protein and 5%–32% lipid by dry mass) can result ...
[146]
Caught in the web: Spider web architecture affects prey ...
May 30, 2018 · Stoichiometry of spider–prey interactions. Web‐building spiders and their prey varied widely in their body nutrient content (Figures 5 and 6) ...
[147]
intraguild prey than pest species in arachnid diets may compromise ...
These web spiders likely consume mainly winged aphids, whereas cursorial spider also consume sedentary life stages, but their role in aphid control may ...
[148]
Toxins for Transgenic Resistance to Hemipteran Pests - PMC
Some continuous feeders such as the leafhoppers Euscelidius variegatus and Eurymela distincta Signoret have a filter chamber to allow excess water from the ...
[149]
EENY016/IN142: Southern Green Stink Bug, Nezara viridula ...
The economic threshold for southern green stink bug in soybeans is reported as 36 stink bugs per 100 swings of a net. For cowpea (southern pea), a population ...
[150]
Brown Marmorated Stink Bug | US EPA
Jun 4, 2025 · They cause major economic damage to fruit, vegetable, and field crops in the mid-Atlantic region.
[151]
Detection of damage caused by Nezara viridula on soybean ... - NIH
The well‐known symptoms of heteropteran damage on harvested soybeans are loss of size, discoloration, patches, and wrinkling. Accompanying the appearance of ...
[152]
[PDF] Soybean Aphids in South Dakota - SDSU Extension
During soybean aphid outbreaks, heavily infested soybean fields may exhibit yield losses of approximately 40%1. As a result of the economic threat posed by ...
[153]
Researchers Helping Protect Crops From Pests - USDA NIFA
Feb 6, 2023 · Aphids and the viruses they transmit damage many food crops, including wheat. ... economic loss will be about $120 million for growers. Hence, the ...
[154]
The Brown Marmorated Stink Bug: Invasive Species Spotlight
Feb 19, 2025 · In 2010, a BMSB outbreak caused over $37 million in losses to fruit producers in this area. These significant economic losses highlight the ...
[155]
[PDF] Economic Impact of the Sugarcane Aphid Outbreak in South Texas
Nov 20, 2017 · Farm profit losses were modeled as changes to the income of households making $75,000–$100,000 annually. Farm losses consisted of revenue losses.
[156]
[PDF] MANAGEMENT OF STINK BUGS (HEMIPTERA - UKnowledge
Dec 10, 2020 · Overall, proper pest management strategies should be incorporated to soybean production to assure sustainable suppression of stink bug insect.
[157]
[PDF] Natural Enemies and Biological Control of Stink Bugs (Hemiptera
Oct 14, 2022 · Stink bugs are historically important agricultural pests in the US, especially in the ... pest, Halyomorpha halys (Hemiptera: Pentatomidae): ...
[158]
[PDF] Economic Threshold for Soybean Aphid (Hemiptera: Aphididae)
The average economic threshold (ET) over all control costs, market values, and yield was 273 ⫾ 38 (mean ⫾ 95% conﬁdence interval [CI], range 111–567) aphids ...
[159]
9. Insect Management | NC State Extension Publications
Jul 31, 2025 · ... insect pests and enhance the soybean crop's ability to tolerate insect injury. ... insect pest of North Carolina soybeans, followed by stink bugs.
[160]
Minute Pirate Bug and Insidious Flower Bug - Cornell CALS
These predatory Orius are beneficial to farmers and gardeners, because they can reduce pest populations.
[161]
Minute pirate bugs - UC IPM
Orius species or other minute pirate bugs are sometimes purchased and released in commercial greenhouses to help control thrips. In most outdoors situations, ...
[162]
Insidious Flower Bug, Minute Pirate Bug Orius insidiosus Say ...
The insidious flower bug, Orius insidiosus Say, is a common predator of a wide variety of small, soft-bodied arthropods.
[163]
Hemiptera - an overview | ScienceDirect Topics
The size of the dorsoventrally flattened insects ranges from 3 to 6 mm. They puncture cutaneous blood vessels of the host with their mandibular stylets ...
[164]
Cimicids of Medical and Veterinary Importance - PMC
Kissing bugs or triatomines are of great medical and veterinary importance because they are vectors of Trypanosoma cruzi, the causative agent of Chagas disease ...
[165]
Chagas Disease, an Endemic Disease in the United States - CDC
Aug 13, 2025 · Triatoma sanguisuga and T. protracta kissing bugs have the largest overall distribution within the United States, but the T. gerstaeckeri ...Missing: 2020s | Show results with:2020s
[166]
DPDx - Bed Bugs - CDC
... disease. The primary medical importance is inflammation associated with their bites (due to allergic reactions to components in their saliva). Cimex lectularius ...Missing: significance | Show results with:significance
[167]
Systemic veterinary drugs for control of the common bed bug, Cimex ...
Nov 17, 2022 · The common bed bug, Cimex lectularius L., is a hematophagous ectoparasite that was a common pest in poultry farms through the 1960s.
[168]
Managing Bed Bugs: A Reemerging Pest in Poultry Production
Mar 6, 2020 · Bed bugs can cause irritating white welts on the skin of chickens and large infestations may lead to excessive feather loss, cloacal irritation, lesions on the ...Missing: veterinary | Show results with:veterinary
[169]
“Kissing Bugs”: Potential Disease Vectors and Cause of Anaphylaxis
Allergic reactions are either localized at the site of the bite with a substantial welt that itches intensely or are systemic with anaphylaxis [20].
[170]
Anaphylactic Reactions Due to Triatoma protracta (Hemiptera ...
The bite from a kissing bug can also lead to serious allergic reactions, including anaphylaxis. A potentially life-threatening allergic response typically needs ...Missing: itch | Show results with:itch
[171]
Bed bugs: clinical relevance and control options - PubMed
Anemia may result when bed bugs are numerous, and their allergens can trigger asthmatic reactions.
[172]
Impact of involving the community in entomological surveillance of ...
Feb 5, 2021 · In Argentina, Triatoma infestans is the principal vector; therefore, vector control is the main strategy for the prevention of this illness. The ...
[173]
Towards environmental detection of Chagas disease vectors and ...
Jun 14, 2022 · Chagas disease vector surveillance typically relies on identification of infested houses using timed-manual collections (TMCs), conducted by ...
[174]
Kissing bug: Chagas disease is now endemic to the US, scientists say
Sep 16, 2025 · Doctors have reported Chagas cases in humans in eight states: Arkansas, Arizona, California, Louisiana, Missouri, Mississippi, Tennessee and ...Missing: 2020s | Show results with:2020s
[175]
Cochineal: A Product of Nature
Cochineal, Dactylopius coccus, is a small scale insect native to subtropical ... This is the key ingredient for producing the highly coveted vibrant red dye.
[176]
(PDF) A review on cochineal (Dactylopius Coccus Costa) dye
Sep 9, 2022 · Cochineal dye, made from the dried bodies of female cochineal insects, has been used for centuries as red colourants for food industry (Deveoglu ...<|separator|>
[177]
Unraveling the Role of Lac Insects in Providing Natural Industrial ...
Dec 5, 2022 · Lac is the only natural raw material derived from insects, being obtained from several species of the genus Kerria. Lac is commercially ...
[178]
Edible and Medicinal Progress of Cryptotympana atrata (Fabricius ...
Oct 5, 2023 · The Cryptotympana atrata is a common resource insect with great economic value in East and South Asia. It has high protein, low fat, low total ...
[179]
Nutritional Composition, Phenolic Compounds and Antioxidant ...
This work evaluates the macronutrients and bioactive compounds of water boatmen eggs obtained from three different locations in Mexico.
[180]
Cicadas In Chinese Art : NPR
May 10, 2021 · Cicadas are actually quite prominent in Chinese literature, art and culture," says Haiyan Lee, a professor of east Asian languages and cultures.
[181]
[PDF] Chapters 1–3: Introduction, cicadas in mythology, art, and literature
or Hemiptera. The élite of this ... We have seen how deeply rooted the cicada is in. Greek mythology and we shall find it no less familiar in folk-lore.
[182]
Protected areas and the future of insect conservation - ScienceDirect
Jun 30, 2022 · Insects are apparently facing similar threats to many other taxa – habitat loss and degradation through deforestation, climate change, resource ...
[183]
Scientists' warning on climate change and insects - ESA Journals
Nov 7, 2022 · With an increase of 3.2°C, the ranges of almost half of the insect species will contract by 50% or more, whereas this drops to 18% of insects at ...
[184]
Cicada as a food for mammals: a global review and implications for ...
Aug 8, 2025 · Almost all cicada species have yet to be registered on the IUCN Red List, but Magicicada spp. were the only Near Threatened species.