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Insect

Insects are a class of hexapod invertebrates within the phylum Arthropoda, subphylum Hexapoda, characterized by a chitinous exoskeleton, three distinct body regions or tagmata (head, thorax, and abdomen), a single pair of antennae, compound eyes (often supplemented by ocelli), three pairs of jointed legs attached to the thorax, and typically one or two pairs of wings arising from the thorax.^[1]^[2]^[3] This body plan enables remarkable adaptability, with insects undergoing incomplete or complete metamorphosis during development, involving molting of the exoskeleton and transformation through larval, pupal (in holometabolous orders), and adult stages.^[2]^[4] With over 1 million described species—representing approximately 80% of all known animal species—insects are the most diverse group of multicellular organisms on Earth, and estimates suggest a total of 5.5 million species may exist, though many remain undiscovered due to their small size and vast habitats.^[5]^[6] They are divided into about 29 orders, with the most species-rich being Coleoptera (beetles, over 350,000 species), Lepidoptera (butterflies and moths, over 180,000 species), and Hymenoptera (ants, bees, and wasps, over 150,000 species).^[2] Insects have evolved over 400 million years, originating in the Devonian period, and now inhabit nearly every terrestrial and freshwater ecosystem, from deserts and rainforests to urban areas, though they are absent from the deep ocean and extreme polar regions.^[4]^[7] Insects play pivotal roles in global ecosystems, serving as primary pollinators for about 75% of flowering plants and major crops, decomposers that recycle nutrients through scavenging and dung burial, predators and parasitoids that regulate pest populations, and a foundational food source for vertebrates and other invertebrates.^[7]^[8] Their activities support biodiversity, soil health, and agriculture, yet some species act as vectors for diseases (e.g., mosquitoes transmitting malaria) or agricultural pests, leading to significant economic impacts estimated in billions annually.^[7] Despite their ecological indispensability, insect populations face widespread declines due to habitat loss, pesticides, and climate change, threatening food security and ecosystem stability.^[9]^[5]

Definition and Basics

Etymology and nomenclature

The word insect derives from the Latin insectum, the neuter past participle of insecare ("to cut into" or "to notch"), reflecting the apparent segmentation of the animal's body into distinct parts.^[10] This term appeared in classical texts, such as Pliny the Elder's Natural History (circa 77 CE), where it translated the Greek entomon ("notched" or "cut up"), but it gained systematic usage in modern taxonomy through Carl Linnaeus's Systema Naturae (10th edition, 1758), in which he established Insecta as a class encompassing arthropods like insects, arachnids, and crustaceans.^[10]^[11] Linnaeus's introduction of binomial nomenclature in the same 1758 edition revolutionized insect naming by assigning each species a unique two-word Latinized identifier: the genus name (capitalized) followed by the specific epithet (lowercase), such as Musca domestica for the housefly.^[11] This system, applied uniformly to insects as part of zoological taxonomy, includes the designation of type species for each genus to anchor nomenclature stability; for example, the type species for the genus Apis (bees) is Apis mellifera, the Western honeybee.^[11] The International Code of Zoological Nomenclature (ICZN), founded in 1895 and revised through its fourth edition in 1999, governs these rules specifically for animals, including insects, addressing revisions like updates to publication requirements and digital naming to maintain nomenclatural consistency.^[12]^[13] Prior to Linnaeus, insect naming lacked standardization, with vernacular terms like "bug" used broadly in English from the late 16th century to denote any small, creepy, or pesty creature, originating from Middle English bugge (possibly from Welsh bwgan, meaning "ghost" or "goblin," evoking fear of the unknown).^[14]^[15] In contrast, modern entomological classifications under the ICZN distinguish scientific binomina from common names; for instance, the "ladybug" is formally Coccinella septempunctata, while "honeybee" corresponds to Apis mellifera.^[13] This shift from folk terminology to precise, universal nomenclature has enabled global collaboration in entomological research.^[12]

Distinguishing features

Insects are distinguished by their tripartite body plan, consisting of a head, thorax, and abdomen, which represents a specialized tagmosis unique among arthropods. The head typically arises from the fusion of six embryonic segments, the thorax comprises three distinct segments (prothorax, mesothorax, and metathorax), and the abdomen usually features eleven segments, though modifications occur across species.^[16]^[17] This segmentation pattern supports specialized functions, with the head housing sensory and feeding structures, the thorax bearing locomotion appendages, and the abdomen dedicated to visceral organs.^[2] A hallmark of insects is their hexapod nature, characterized by exactly three pairs of jointed legs attached to the thoracic segments, setting them apart from other arthropods such as arachnids with eight legs or crustaceans with variable limb counts exceeding six.^[18]^[19] These legs, each composed of segments including coxa, trochanter, femur, tibia, tarsus, and pretarsus, enable diverse adaptations like running, jumping, or grasping.^[17] Many adult insects also possess one or two pairs of wings emerging from the meso- and metathorax, facilitating flight and dispersal, though wingless forms exist in primitive or specialized groups.^[4]^[20] Sensory structures further define insects, including paired compound eyes on the head that provide panoramic vision through thousands of ommatidia and one pair of antennae serving as primary chemoreceptors and mechanosensors.^[2]^[21] The entire body is encased in a chitinous exoskeleton, or integument, which offers protection, structural support, and sites for muscle attachment while preventing desiccation.^[4]^[20] Physiologically, insects exhibit an open circulatory system where hemolymph bathes tissues directly, pumped by a dorsal vessel acting as a heart, contrasting with the closed systems of vertebrates.^[2]^[4] Respiration occurs via a tracheal system of branching tubes that deliver oxygen directly to cells, entering through spiracles on the thorax and abdomen, an efficient adaptation for their small size and high metabolic demands.^[2]^[20] These traits collectively underscore the class Insecta's evolutionary specialization within the phylum Arthropoda.^[19]

Diversity and distribution

Insects represent one of the most species-rich groups on Earth, with approximately 1 million species formally described and estimates suggesting a total of 5 to 10 million species worldwide.^[22] Among the described species, the order Coleoptera (beetles) exhibits the highest diversity, with over 400,000 species, while Lepidoptera (butterflies and moths) follows with approximately 180,000 species.^[23]^[24] These figures underscore the immense undescribed biodiversity, particularly in understudied tropical regions where new species discoveries continue at a steady pace. Insects are distributed across nearly all terrestrial and freshwater habitats globally, achieving ubiquity except in the deep ocean and polar ice caps, where extreme conditions preclude their presence.^[25] Their highest densities and species richness occur in tropical rainforests, which harbor the majority of global insect diversity due to stable climates and abundant resources fostering speciation.^[26] Biogeographic patterns reveal pronounced endemism in isolated regions; for instance, Madagascar's insect fauna includes a high proportion of unique species, with over 90% endemism in well-studied groups like ants and mayflies, resulting from the island's long geological isolation.^[27]^[28] Diversity patterns are shaped by environmental factors, including habitat fragmentation, which can reduce local species richness by isolating populations and limiting gene flow, while climate influences speciation rates through variations in temperature and precipitation that drive adaptive radiations.^[29]^[30] These dynamics highlight how connectivity and climatic stability in biodiverse hotspots like the tropics promote evolutionary divergence, contributing to the overall global mosaic of insect distributions.

Evolutionary Origins

Phylogenetic relationships

Insects belong to the phylum Arthropoda, where they are classified within the subphylum Hexapoda, encompassing all six-limbed arthropods including true insects and their close relatives such as springtails and proturans.^[31] Within Arthropoda, Hexapoda forms the clade Pancrustacea alongside Crustacea, with molecular phylogenomic analyses of nuclear protein-coding sequences strongly supporting Hexapoda as the sister group to a derived lineage of crustaceans known as Xenocarida (comprising remipeds, cephalocarids, and branchiopods). Recent phylogenomic studies using extensive genomic data continue to affirm this relationship.^[31]^[32] This relationship positions insects as nested within a paraphyletic Crustacea, rejecting earlier hypotheses that linked insects more closely to myriapods (centipedes and millipedes).^[31] Broader arthropod phylogeny places the Pancrustacea clade as sister to Myriapoda within the Mandibulata, a grouping defined by shared mandibular mouthparts, while Chelicerata (including spiders, scorpions, and horseshoe crabs) branches basally from Mandibulata, supported by extensive phylogenomic data from over 60 nuclear genes across 178 arthropod species.^[31] Arthropoda as a whole resides within the superphylum Ecdysozoa, a molting clade of protostome bilaterians that also includes nematodes and onychophorans, distinguishing arthropods from lophotrochozoan bilaterians such as annelids and mollusks through molecular markers like 18S ribosomal RNA sequences.^[33] Molecular evidence robustly affirms the monophyly of Insecta (or more broadly Hexapoda), with analyses of 18S rRNA genes from multiple arthropod taxa revealing shared derived nucleotide substitutions that unite hexapods exclusive of other arthropods, as demonstrated in combined datasets of ribosomal and protein-coding loci.^[33] Similarly, Hox gene cluster comparisons across arthropods, including sequences from basal hexapods like silverfish, support hexapod monophyly through conserved genomic organization and expression patterns, such as the duplication and divergence of the Antennapedia and fushi tarazu genes specific to insects.^[34] Within Hexapoda, the major clades contrast the wingless Apterygota—comprising Archaeognatha (bristletails) and Zygentoma (silverfish and firebrats), which form a paraphyletic grade of primitive, ametabolous insects—with the monophyletic Pterygota, encompassing all winged insects and secondarily wingless forms like fleas.^[35] Mitochondrial genome phylogenies from basal lineages confirm Zygentoma as the immediate sister group to Pterygota (forming the Dicondylia), while Archaeognatha diverges earlier, underscoring the evolutionary transition from wingless ancestors to the dominant winged radiation.^[35] This division highlights the origin of flight as a key innovation within Pterygota, with fossil evidence from the Devonian period aligning with molecular divergence estimates around 400 million years ago.^[35]

Fossil record and timeline

The fossil record of insects begins in the Devonian period, approximately 407 to 396 million years ago, with the fragmentary remains of Rhyniognatha hirsti from the Rhynie Chert in Scotland, representing a potential early insect based on its dicondylic mandibles suggestive of a pterygote affinity.^[36] This specimen, consisting primarily of preserved mouthparts, indicates that insects may have originated near the base of the Ectognatha clade, though its exact placement remains debated due to limited material and possible affinities with other arthropods.^[36] Subsequent Devonian evidence is scarce, highlighting a significant "Hexapod Gap" spanning much of this period to the early Carboniferous, during which few definitive insect traces appear, likely reflecting preservational biases rather than absence.^[37] The Carboniferous period, particularly its later stages around 328 to 324 million years ago, marks a major diversification of winged insects (Pterygota), with fossils documenting the rapid emergence of diverse orders such as Palaeodictyopteroidea and early odonatans featuring large, veined wings adapted for flight.^[37] This radiation coincided with expanding terrestrial forests and elevated atmospheric oxygen levels, enabling larger body sizes and aerial capabilities, as evidenced by compressions and impressions from coal-bearing deposits. By the Pennsylvanian subperiod (roughly 323 to 299 million years ago), winged forms dominated the record, with over 1,000 described species illustrating a burst in morphological disparity. In the Mesozoic era, insect diversification accelerated further, particularly during the Cretaceous period (145 to 66 million years ago), aligning with the radiation of angiosperms that provided new ecological niches through floral resources and pollination mutualisms.^[38] This era saw the proliferation of holometabolous insects, including bees, butterflies, and beetles, with family-level diversity stabilizing at modern levels by the mid-Cretaceous, driven by angiosperm expansion that mitigated extinctions and boosted originations.^[38] Key evidence comes from lagerstätten preserving soft tissues, such as the Mazon Creek deposits in Illinois (late Carboniferous, ~308 million years ago), which yield exceptionally preserved insects like orthopterans and palaeodictyopterans in siderite concretions, offering insights into early terrestrial ecosystems.^[39] Similarly, the Crato Formation in Brazil's Araripe Basin (Aptian stage, ~113 million years ago) has produced over 2,000 insect specimens, including odonates and hemipterans with mineralized cuticles, revealing high-fidelity taphonomic windows into Cretaceous biodiversity.^[40] Despite these rich assemblages, the insect fossil record remains highly incomplete, primarily due to the small size, soft-bodied nature, and terrestrial habits of most species, which favor rapid decay over mineralization.^[37] Estimates suggest that the documented fossil diversity captures only a fraction of past insect richness, as indicated by sampling biases in lagerstätten and the underrepresentation of immature stages or non-aquatic forms.^[41] This incompleteness underscores the need for continued exploration of exceptional deposits to refine timelines of insect evolution.

Key evolutionary adaptations

Insects achieved remarkable success through several pivotal evolutionary innovations that enhanced their survival, dispersal, and diversification. One of the most transformative adaptations was the development of powered flight in the Pterygota clade, which originated approximately 350 million years ago during the early Carboniferous period.^[42] This innovation predated vertebrate flight by approximately 130 million years and enabled insects to escape predators, overcome geographic barriers for migration, access new food resources, and locate mates more effectively, contributing to their rapid diversification into over 15 orders by the end of the Carboniferous.^[42] Another critical adaptation was the evolution of metamorphosis, particularly the complete (holometabolous) form, which emerged around 350 million years ago in the early Carboniferous.^[43] This developmental strategy involves distinct larval, pupal, and adult stages, allowing for niche separation where larvae specialize in feeding and growth in protected or aquatic environments, while adults focus on reproduction and dispersal in often contrasting habitats.^[44] The hormonal regulation, involving ecdysteroids for molting and juvenile hormones for stage specification, facilitated this separation, reducing intraspecific competition and enhancing overall ecological adaptability.^[43] Holometaboly, seen in over 60% of insect species today, such as beetles and butterflies, underscores its role in driving insect dominance.^[44] Refinements to the exoskeleton, particularly the incorporation of cuticular hydrocarbons (CHCs) into the chitin-based cuticle, were essential for resisting desiccation and conquering terrestrial environments. This waterproofing layer evolved as arthropods, including early insects, transitioned to land around 400 million years ago, with CHCs forming a lipid barrier that minimizes water loss through the epicuticle.^[45] In Drosophila species, variations in CHC chain length and branching explain much of the differences in desiccation tolerance, with longer methyl-branched CHCs in arid-adapted lineages enhancing survival in dry conditions.^[45] These modifications, building on the basic chitin-protein structure, provided mechanical support and protection while enabling insects to thrive in diverse terrestrial habitats without constant access to water.^[46] The co-evolution of insects with angiosperms following the Cretaceous period further propelled insect diversification through specialized herbivory and pollination interactions. Angiosperm radiation during the Early Cretaceous (125–90 million years ago) initiated the Cretaceous Terrestrial Revolution, but post-Cretaceous dynamics, particularly the Angiosperm Terrestrial Revolution (100–50 million years ago), amplified insect origination rates while mitigating extinctions.^[38] Fossil evidence from around 99 million years ago shows early pollination syndromes, with insects like bees and butterflies developing morphological and behavioral specializations for angiosperm pollen transfer, while herbivorous clades adapted to exploit new plant tissues and defenses.^[38] This reciprocal evolution, evident in orders such as Lepidoptera and Diptera, linked insect family richness peaks to angiosperm dominance, fostering biodiversity through mutual dependencies.^[38]

Physical Structure

External morphology

The external morphology of insects is characterized by a chitinous exoskeleton that provides support and protection, divided into three main tagmata: the head, thorax, and abdomen.^[47] This exoskeleton, known as the cuticle, consists of multiple layers and is periodically shed during molting to allow for growth.^[48] The head is a hardened capsule formed by sclerites, which are chitinous plates fused together, enclosing sensory organs and mouthparts.^[47] Insect mouthparts exhibit diverse modifications adapted to feeding habits; for instance, chewing mouthparts feature laterally moving mandibles for biting and grinding solid food, as seen in grasshoppers, while piercing-sucking types involve stylets that penetrate tissues to extract liquids, typical in aphids.^[49]^[47] Antennae, paired appendages on the head, vary in shape for sensory functions; filiform antennae are thread-like with uniform segments, common in ground beetles, whereas clubbed forms widen distally, as in butterflies with capitate or clavate types.^[49]^[47] The thorax comprises three segments—the prothorax, mesothorax, and metathorax—each bearing a pair of legs and, in pterygotes, wings attached to the meso- and metathorax.^[49]^[47] Legs are segmented appendages consisting of the coxa (basal segment attached to the thorax), trochanter, femur (largest segment), tibia, and tarsus (distal foot-like portion, often with claws).^[47] Wings, when present, are typically membranous with intricate venation patterns formed by chitinous veins that reinforce the structure and aid in species identification; for example, dragonflies display extensive branching veins, while butterflies have reduced venation in scaled wings.^[49]^[47] The abdomen is a flexible region typically composed of 11 segments, each with dorsal terga and ventral sterna sclerites connected by intersegmental membranes for expansion during feeding or reproduction.^[47] In many species, the terminal segments bear cerci, sensory appendages that detect vibrations or chemicals, and females often possess an ovipositor, a specialized structure for depositing eggs into substrates.^[49]^[47] The cuticle, the outermost layer enveloping the body, is secreted by the underlying epidermal cells and divided into the epicuticle and procuticle.^[48] The epicuticle is a thin, waxy outer coating with sublayers including a cement layer, wax layer, and inner epicuticle, primarily functioning to prevent desiccation; studies show variations in wax composition influence water loss rates across species and habitats.^[48] The procuticle, beneath the epicuticle, comprises an exocuticle (sclerotized for rigidity via protein tanning) and endocuticle (flexible, containing chitin microfibrils in a protein matrix), providing mechanical strength.^[48] Molting, or ecdysis, occurs when the old cuticle splits along predetermined lines, allowing the insect to emerge and expand a new, soft cuticle that hardens over hours to days, enabling growth in a rigid exoskeleton.^[48]^[47]

Internal organ systems

Insects possess a diverse array of internal organ systems adapted for efficient physiological function within their exoskeleton-constrained bodies. These systems facilitate essential processes such as nutrient processing, gas exchange, circulation, neural coordination, and reproduction, often differing markedly from those in vertebrates due to the insects' open circulatory design and segmented architecture. The nervous system of insects is decentralized and comprises a dorsal brain located in the head, a ventral nerve cord running along the ventral body surface, and segmental ganglia that serve as local processing centers. The brain integrates sensory inputs and coordinates complex behaviors, while the ventral nerve cord connects the brain to fused thoracic and abdominal ganglia, allowing for rapid reflex responses in each body segment. This structure enables efficient control over locomotion and environmental interactions without a centralized spinal cord.^[50]^[51]^[52] Visual processing is handled by compound eyes and ocelli, which interface directly with the nervous system. Compound eyes consist of numerous ommatidia, each functioning as an independent optical unit with a corneal lens, crystalline cone, and photoreceptor cells that detect light via rhabdomeric phototransduction, providing wide-angle vision suited for motion detection. Ocelli, simpler dorsal eyes with a single lens and fewer photoreceptors, primarily sense light intensity and horizon orientation to aid in flight stabilization.^[53]^[54] The digestive system forms a complete tubular alimentary canal divided into foregut, midgut, and hindgut regions, each with specialized roles in ingestion, nutrient absorption, and waste elimination. The foregut, lined with cuticle, includes the mouth, pharynx, esophagus, crop for storage, and proventriculus for grinding; the midgut is the primary site of enzymatic digestion and absorption, often protected by a peritrophic membrane; and the hindgut reabsorbs water and ions from wastes before expulsion via the anus. Malpighian tubules, blind-ending structures arising at the midgut-hindgut junction, function in excretion by filtering hemolymph to remove nitrogenous wastes like uric acid, which are then processed in the hindgut for efficient water conservation in terrestrial environments.^[55]^[56]^[57] Circulation relies on an open system where nutrient-rich hemolymph bathes tissues directly within the hemocoel cavity, pumped by a dorsal vessel that acts as both heart and aorta. The posterior abdominal portion functions as the heart, with ostia (valved openings) allowing hemolymph entry during diastole, while muscular contractions propel it anteriorly through the vessel; the anterior thoracic aorta distributes it forward before it diffuses back. Hemolymph, comprising plasma, hemocytes for immunity, and minimal respiratory pigments, lacks hemoglobin and relies on body movements and accessory pumps for circulation, ensuring oxygen and nutrient delivery despite low pressure.^[58]^[59]^[60] Respiration occurs via a tracheal system of air-filled tubes that deliver oxygen directly to cells, bypassing blood transport. External spiracles on the thorax and abdomen open into tracheae, which branch into finer tracheoles penetrating tissues; gas exchange happens by diffusion across thin tracheole walls, driven by concentration gradients and enhanced in active insects by abdominal pumping or spiracle regulation. This system supports high metabolic rates during flight, with spiracles often valved to minimize water loss, achieving efficient O₂ uptake comparable to vertebrate lungs in small-bodied insects.^[61]^[62]^[63] The reproductive system includes paired ovaries or testes in the abdomen, producing gametes, along with accessory glands that secrete supportive fluids. In females, ovaries consist of ovarioles where oocytes develop, maturing into eggs released via oviducts to a genital chamber; accessory glands produce yolk proteins, adhesives, or protective coatings. Males have testes forming spermatocytes that mature into sperm stored in seminal vesicles, transferred via ejaculatory ducts with contributions from accessory glands for spermatophore formation. Endocrine regulation involves ecdysone, a steroid hormone from prothoracic glands that triggers vitellogenesis and egg maturation, and juvenile hormone from corpora allata, which modulates reproductive development and prevents premature metamorphosis in adults.^[64]^[65]^[66]

Reproduction and Growth

Mating and fertilization

While the majority of insects reproduce sexually, some species are capable of asexual reproduction through parthenogenesis, in which unfertilized eggs develop into offspring, typically females. This mode is common in aphids, which use cyclical parthenogenesis to rapidly increase populations during favorable conditions, and in certain stick insects (Phasmatodea), where females produce all-female broods without males.^[67] Insects exhibit diverse mating behaviors that ensure successful reproduction, primarily involving internal fertilization where sperm is transferred from males to females during copulation.^[68] Courtship rituals play a crucial role in mate attraction and selection, often relying on chemical, visual, or auditory signals to synchronize mating. These behaviors vary widely across species, reflecting adaptations to environmental and ecological pressures.^[69] Courtship in many insects begins with pheromones, volatile chemicals released by one sex to attract the opposite sex over distances. For instance, female moths release sex pheromones that guide males to potential mates using olfactory cues. In honey bees (Apis mellifera), virgin queens produce pheromones during nuptial flights that draw drones to mating congregations, facilitating airborne copulation.^[70] Visual and behavioral displays complement these signals; male fireflies (Photinus species) emit species-specific bioluminescent flashes in patterned sequences to court females, with receptive females responding via flashes to indicate acceptance.^[71] Similarly, male fruit flies (Drosophila melanogaster) perform elaborate dances involving wing vibrations and leg taps to stimulate females during courtship.^[72] Insect mating systems range from monogamy to polygamy, though polygamy—where individuals mate with multiple partners—is predominant, allowing for higher reproductive output in resource-limited environments.^[73] Monogamy occurs rarely, often in species with high paternal investment like certain burying beetles, but most insects, such as butterflies and cockroaches, engage in promiscuous mating to maximize genetic diversity in offspring.^[74] Sperm transfer typically occurs via direct insemination through male genitalia or indirectly via a spermatophore, a proteinaceous packet containing sperm and nutrients produced by male accessory glands. In lepidopterans like moths, the spermatophore is deposited in the female's reproductive tract during copulation, providing both genetic material and sustenance to enhance egg production.^[75] Fertilization is invariably internal in insects, with sperm stored in female spermathecae for delayed use in egg fertilization. A notable variation is traumatic insemination, observed in bed bugs (Cimex lectularius), where males pierce the female's abdominal wall with a specialized paramere to inject sperm directly into the hemocoel, bypassing the genital tract.^[76] This coercive strategy incurs costs to females, including injury and immune response, but has led to counter-adaptations like the spermalege, a specialized organ that reduces damage.^[77] Variations in parental investment during mating include short-term mate guarding and nuptial gifts, which provide immediate benefits to females without extending to prolonged offspring care. In some bush crickets, males transfer spermatophores containing nutritious lipids that females digest to boost fecundity, representing a form of paternal investment that influences female remating decisions.^[78] Males in species like the green lacewing may guard females post-copulation to prevent sperm competition from rival males, ensuring higher paternity success.^[79] These strategies highlight the evolutionary trade-offs in insect reproduction, balancing male reproductive assurance with female control over fertilization.^[80]

Developmental stages and metamorphosis

Insects exhibit a remarkable diversity in their developmental processes, primarily categorized into three types of metamorphosis: ametabolous, hemimetabolous, and holometabolous. These variations reflect evolutionary adaptations in life cycle strategies, allowing insects to occupy diverse ecological niches. Ametabolous development represents the most primitive form, with no distinct metamorphic stages, while hemimetabolous and holometabolous forms involve increasing degrees of transformation between juvenile and adult phases.^[81] Ametabolous development, seen in primitive orders such as Zygentoma (e.g., silverfish), involves direct growth without significant morphological changes between juveniles and adults. Juveniles hatch from eggs resembling miniature adults and undergo multiple molts throughout life, with adults continuing to molt periodically to replace worn exoskeletons. This pattern lacks a pupal stage or major restructuring, emphasizing continuous growth rather than transformation.^[81] Hemimetabolous, or incomplete, metamorphosis occurs in orders like Orthoptera (e.g., grasshoppers) and Hemiptera (e.g., true bugs), featuring egg, nymph, and adult stages. Nymphs emerge from eggs and closely resemble adults in form and habitat but are wingless and sexually immature initially. Through successive molts—typically 4 to 8 instars—nymphs gradually develop wings, genitalia, and other adult features, with each instar becoming progressively more adult-like. The final molt produces a fully winged, reproductive adult, without a quiescent pupal phase.^[81] Holometabolous, or complete, metamorphosis is the most derived and widespread pattern, found in over 80% of insect species, including Coleoptera (beetles) and Lepidoptera (butterflies). It encompasses four distinct stages: egg, larva, pupa, and adult. Larvae hatch as worm-like, feeding specialists often dissimilar to adults, undergoing several molts to grow while remaining in the larval form. The prepupal molt leads to the pupal stage, a non-feeding, immobile phase where extensive tissue remodeling occurs—larval structures histolyze, and adult organs differentiate from imaginal discs. The adult ecloses from the pupa fully formed, with wings, reproductive organs, and other mature features. For instance, in butterflies like Danaus plexippus (monarch), the caterpillar larva feeds voraciously before pupating into a chrysalis, emerging as a winged adult capable of migration. This separation of feeding (larval) and reproductive (adult) phases enhances resource partitioning.^[81] These metamorphic processes are tightly regulated by hormones, primarily ecdysone and juvenile hormone (JH). Ecdysone, a steroid hormone produced by the prothoracic glands, initiates molting and metamorphic changes by triggering gene expression cascades that lead to apolysis (exoskeleton separation) and new cuticle formation. In all insect types, pulses of ecdysone drive each molt. JH, a sesquiterpenoid secreted by the corpora allata, modulates the response to ecdysone: high levels maintain juvenile characteristics and prevent premature metamorphosis, while declining levels allow ecdysone to promote adult differentiation. In ametabolous and hemimetabolous insects, JH persists through most molts but drops before the final adult molt; in holometabolous forms, JH is low during the larval-to-pupal transition, enabling complete restructuring. Disruptions in this hormonal balance, such as altered JH titers, can lead to developmental anomalies like extra larval instars or failed ecdysis.^[81]^[82]

Behavioral Patterns

Communication methods

Insects employ a diverse array of communication methods to interact with conspecifics, including chemical, visual, auditory, and tactile signals, which facilitate coordination in mating, foraging, and social behaviors. These modalities are adapted to the insects' sensory capabilities and environmental constraints, often overlapping in function to enhance signal reliability. Pheromones represent the primary chemical communication channel in insects, consisting of volatile or contact semiochemicals that elicit specific behavioral or physiological responses. Alarm pheromones, such as those released by aphids or ants during threats, trigger rapid escape or defensive aggregation among nearby individuals. Sex pheromones, crucial for mate attraction, are exemplified by bombykol (E,Z)-10,12-hexadecadien-1-ol, a 16-carbon alcohol produced by female silkworms (Bombyx mori) that stimulates males to initiate courtship flights over long distances. Trail pheromones, laid by social insects like ants, guide colony members to food sources; for instance, the Argentine ant (Linepithema humile) uses a blend of hydrocarbons to mark persistent foraging paths. These pheromones are detected via specialized antennal sensilla, with structures varying by type—e.g., bombykol binds to odorant receptors in moth antennae to activate neural signaling. Visual signals in insects often rely on color patterns and movements visible under daylight or low-light conditions, serving to attract mates or deter rivals. Wing patterns, such as the iridescent scales on butterfly wings or the eyespots on moth hindwings, convey species identity or warning signals during courtship displays. Bioluminescence provides a striking visual cue in certain beetles, notably fireflies (family Lampyridae), where light is produced through the oxidation of luciferin catalyzed by luciferase in photocytes, emitting flashes in species-specific patterns to synchronize mating. This reaction yields a cold light peaking at around 560 nm, efficient for nocturnal communication without significant heat loss.^[83] Auditory communication involves the production and detection of airborne sounds or substrate-borne vibrations, enabling interactions over short to medium ranges. Stridulation, a common mechanism, occurs when body parts are rubbed together; in crickets (Gryllidae), males rub their forewings to produce chirps via file-and-scraper structures, with the song's frequency and pulse rate signaling fitness to females. Substrate vibrations, transmitted through plants or soil, are used by insects like leafhoppers, where tymbal organs generate pulses detected by subgenual organs in the legs, facilitating mate location or aggregation. These signals are particularly effective in dense vegetation where visual cues are obscured. Tactile communication manifests through direct physical contact, often involving antennae or body grooming to exchange information in close proximity. Antennation, where insects touch antennae to one another, allows social species like bees to transfer pheromonal cues or assess colony status, as seen in honeybees (Apis mellifera) during trophallaxis. Grooming behaviors, such as mutual antennal cleaning in termites, reinforce social bonds and distribute alarm signals within nests. These methods are integral to maintaining cohesion in eusocial groups, though they also play roles in broader interactions like mating recognition. Insects exhibit a range of social structures, from solitary living to highly organized colonial societies, with eusociality representing the most complex form observed primarily in the orders Hymenoptera (ants, bees, and wasps) and Isoptera (termites). Eusocial species are characterized by cooperative brood care, overlapping generations, and a reproductive division of labor where most individuals forgo personal reproduction to support the colony.^[84] In these societies, distinct castes emerge, including queens or kings dedicated to reproduction, sterile workers focused on foraging, nest maintenance, and brood rearing, and soldiers specialized for defense against intruders.^[85] For instance, in termite colonies, soldiers possess enlarged mandibles for combat, while in ants and bees, workers often display morphological adaptations like reduced wings or enhanced sensory organs suited to their tasks.^[86] The evolution of altruism in eusocial insects, where non-reproductive castes sacrifice their fitness to benefit relatives, is explained by kin selection theory, as formalized by W.D. Hamilton. This theory posits that such behaviors spread if the genetic relatedness (r) between altruist and beneficiary, multiplied by the fitness benefit (B) to the beneficiary, exceeds the fitness cost (C) to the altruist, expressed as Hamilton's rule: rB > C. In Hymenoptera, haplodiploid sex determination results in sisters sharing 75% of their genes on average, elevating r and favoring worker sterility to promote queens' offspring production.^[87] This mechanism underpins the stability of castes, as workers gain indirect fitness through aiding close kin rather than reproducing themselves.^[88] While many insects are solitary, relying on individual efforts for survival and reproduction without cooperative interactions, colonial species like honeybees (Apis mellifera) demonstrate advanced eusocial organization through temporal division of labor. In honeybee hives, workers progress from in-hive duties such as nursing larvae and cleaning to foraging outside as they age, optimizing colony efficiency and resource allocation.^[89] This contrasts with solitary bees, which lack castes and perform all tasks independently, highlighting how eusociality enhances colony resilience against environmental pressures.^[90] Beyond eusociality, non-eusocial insects form temporary aggregations for mutual benefit, as seen in bark beetles (family Curculionidae, subfamily Scolytinae). These beetles, which are otherwise solitary, release aggregation pheromones during host tree colonization, drawing conspecifics to amplify attack success on defended conifers by overwhelming tree defenses through mass infestation.^[91] Such groupings facilitate mating and resource exploitation without permanent castes or reproductive suppression, relying instead on chemical signals to coordinate transient assemblies.^[92]

Modes of locomotion

Insects employ diverse modes of locomotion adapted to terrestrial, aerial, and aquatic environments, leveraging specialized anatomical features for efficient movement. These include walking on varied surfaces, jumping for escape or predation, swimming through water via propulsion mechanisms, and flight powered by oscillatory wing motions. Each mode relies on integrated musculoskeletal systems that enable rapid, energy-efficient travel across scales from millimeters to meters. Flight in insects is primarily achieved through the rapid oscillation of wings, driven by two main muscle types: synchronous and asynchronous. In synchronous flight muscles, each contraction is directly triggered by a neural impulse, limiting wingbeat frequencies to 5–50 Hz in larger insects like butterflies and locusts.^[93] Asynchronous muscles, prevalent in flies and bees, operate via stretch-activation where a single neural input initiates multiple contractions through mechanical feedback, enabling wingbeat frequencies exceeding 100 Hz and up to 200 Hz in species such as Drosophila melanogaster. For instance, fruit fly motor neurons fire at approximately 5 Hz, yet asynchronous muscles generate 200 Hz oscillations for sustained flight. Wing morphology, with flexible hinges and varying vein patterns, further optimizes aerodynamic forces during these beats. Walking is the most common terrestrial locomotion in insects, often utilizing an alternating tripod gait for stability and speed. In this pattern, three legs—one foreleg and hindleg from one side, plus the middle leg from the opposite side—remain in contact with the substrate while the other three swing forward, maintaining balance even on uneven terrain. Cockroaches exemplify this gait, transitioning from slow ambling to fast trotting while sustaining the tripod coordination up to speeds of 1.5 body lengths per second. For vertical climbing on smooth surfaces, many insects rely on adhesive setae—microscopic, hair-like structures on their tarsi that exploit van der Waals forces for attachment. These setae, angled and compliant in flies and beetles, allow reversible adhesion without residue, supporting body weights on ceilings or walls. Aquatic locomotion varies by species, with surface-dwelling insects like water striders (Gerridae) using hydrofuge hairs to exploit surface tension. These hydrophobic microhairs, numbering thousands per square millimeter on legs and body, trap air and repel water, enabling the insects to stride across water films at speeds up to 1.5 meters per second without breaking the surface. In submerged environments, certain larvae employ jet propulsion for rapid movement. Dragonfly larvae (Anisoptera), for example, fill a rectal chamber with water and expel it forcefully through the anus at frequencies up to 2.2 cycles per second, achieving bursts of acceleration for predation or evasion. Jumping serves as a burst locomotion mode for escaping threats or capturing prey, often powered by elastic energy storage. Fleas (Siphonaptera) utilize a catapult mechanism in their hind legs, where the trochanteral extensor muscle compresses a resilin pad—a rubber-like protein with high elasticity—storing energy before rapid release. This enables jumps up to 150 times body length, with launch accelerations exceeding 100 g, far surpassing direct muscle-powered leaps.

Ecological Roles

Habitats and environmental adaptations

Insects predominantly occupy terrestrial habitats, where their exoskeleton serves as a primary barrier against desiccation, particularly in arid environments. The cuticle is coated with hydrocarbons and waxes that form a hydrophobic layer, drastically reducing evaporative water loss through transpiration. In desert-dwelling tenebrionid beetles, such as Onymacris plana, this cuticular permeability is exceptionally low, enabling the insects to prioritize hemolymph dehydration over vital tissue loss during prolonged exposure to dry conditions.^[94] Similarly, species like Rhytinota praelonga regulate body fluid volumes effectively in hyper-arid zones through enhanced wax deposition, which correlates with their survival in environments where relative humidity often falls below 20%.^[94] These adaptations underscore the terrestrial dominance of insects, with over 90% of known species restricted to land-based ecosystems despite their ancient aquatic origins.^[95] Aquatic habitats host a diverse array of insects, especially in larval stages, supported by specialized respiratory mechanisms to extract oxygen from water. Dragonfly nymphs (Odonata: Anisoptera) feature internal gill-like structures within the rectal chamber, where jet-propelled water currents facilitate passive diffusion of dissolved oxygen across thin epithelial linings.^[96] This adaptation not only sustains respiration but also aids in rapid escape from predators via abdominal pumping. Predaceous diving beetles (Coleoptera: Dytiscidae), in contrast, rely on air bubbles trapped beneath their elytra or by hydrofuge hairs on the ventral surface, creating a "physical gill" that allows oxygen replenishment from surrounding water until the bubble diminishes due to nitrogen diffusion.^[96] Such innovations enable these insects to exploit freshwater systems ranging from stagnant ponds to fast-flowing streams, where oxygen levels vary widely.^[96] Insects exhibit profound thermal and altitudinal tolerances, often through physiological dormancy and biochemical defenses against cold stress. Diapause, a hormonally induced arrest in development, allows overwintering by suppressing metabolism, ceasing feeding, and directing energy toward protective sites like soil or plant litter; for example, the European corn borer (Ostrinia nubilalis) enters diapause as a fifth-instar larva to endure subzero temperatures.^[97] Freeze-tolerant species, such as the goldenrod gall fly (Eurosta solidaginis), accumulate cryoprotectants like glycerol and antifreeze proteins—comprising up to 30% of body weight—to nucleate extracellular ice while preserving supercooling in intracellular fluids, preventing lethal crystal damage at temperatures as low as -40°C.^[97] These mechanisms facilitate colonization of high-altitude montane zones and polar fringes, where seasonal extremes challenge survival.^[97] Urban and extreme anthropogenic habitats have become refugia for resilient insect species, particularly invasive peridomestic pests. Cockroaches (Blattodea), such as the German cockroach (Blattella germanica), thrive in human structures by exploiting warm, humid microenvironments like kitchen crevices and basements, where they navigate cracks as narrow as 1/16 inch and forage nocturnally on organic debris.^[98] Their rapid reproduction—yielding over 30,000 offspring per female annually—and tolerance to fluctuating temperatures and low humidity enable persistence in diverse built environments, from apartments to hospitals.^[98] This adaptability highlights insects' capacity to invade novel, resource-rich niches altered by human activity.^[98]

Interactions in ecosystems

Insects serve as pivotal agents in ecosystem dynamics, particularly through their roles in pollination, where they facilitate the reproduction of numerous plant species via mutualistic interactions. Approximately 85–90% of the world's angiosperms depend on animal pollinators, with insects such as bees comprising the majority of these vectors, transferring pollen between flowers in exchange for nectar and pollen rewards.^[99] This process underpins food webs by enabling seed and fruit production, supporting higher trophic levels. However, ongoing insect population declines, driven by habitat loss and climate change as of 2025, are reducing pollination services and threatening plant diversity.^[100] Coevolutionary mutualisms between insects and plants have shaped biodiversity, with specialized floral traits evolving alongside pollinator behaviors over millions of years, as evidenced in tropical communities where functional specialization is pronounced.^[101]^[102]^[102] Decomposition by insects accelerates nutrient cycling, transforming organic detritus into forms accessible to plants and microorganisms. Ants and termites together, particularly in tropical ecosystems, represent about one-third of animal biomass and drive bioturbation, mobilizing carbon, nitrogen, and phosphorus through the breakdown of wood and litter, thereby enhancing soil fertility.^[103] Flies, including blowflies and flesh flies, act as primary colonizers of carrion, with their larvae rapidly consuming tissues and releasing nutrients into the soil, which alters local pH and nutrient concentrations to support microbial activity and plant growth. Overall, insect-mediated decomposition can increase rates by up to 44% in tropical rainforests, preventing nutrient lockup and maintaining ecosystem productivity.^[103]^[103]^[104] In food webs, insects occupy diverse trophic positions, with herbivory and predation influencing population dynamics across levels. Herbivorous insects like aphids function as primary consumers, feeding on plant sap and exerting selective pressure on vegetation, often acting as pests that reduce crop yields while channeling energy upward. Predators such as ladybird beetles (ladybugs) occupy higher trophic levels as secondary consumers, effectively controlling aphid populations by consuming over 50% in controlled settings, thereby stabilizing herbivore outbreaks and preserving plant biomass. These interactions exemplify top-down regulation, where predators mitigate herbivory impacts, fostering balanced energy flow from producers to consumers.^[105]^[105] Symbiotic relationships further amplify insects' ecosystem contributions, notably in termites where gut microbes enable the digestion of recalcitrant cellulose. In wood-feeding termites like Nasutitermes species, diverse bacterial communities—including Firmicutes such as Clostridium termitidis and Spirochaetes—produce lignocellulolytic enzymes that break down plant cell walls, converting up to 45% of substrates like wheat straw into bioavailable carboxylates such as acetate. This mutualism allows termites to access nutrients from lignocellulose, recycling them into soil via frass, and highlights the prokaryotic microbiome's role in sustaining detrital pathways.^[106]^[106]

Defense mechanisms

Insects employ a diverse array of defense mechanisms to deter predators and evade threats, enhancing their survival in competitive environments. These strategies range from physical concealment and behavioral adaptations to chemical warfare and visual warnings, often evolving in response to specific ecological pressures. Such mechanisms not only protect individuals but also contribute to the resilience of insect populations across taxa. Cryptic camouflage allows insects to blend seamlessly into their surroundings, reducing detection by predators. For instance, many katydids exhibit leaf mimicry, where their body morphology and coloration imitate foliage, including veins and edges, to avoid visual predation. This adaptation is particularly effective in diurnal species, as evidenced by studies on Panacanthus species in neotropical forests. Countershading, another form of crypsis, involves darker dorsal surfaces and lighter ventral ones, creating a flattened appearance against backgrounds and minimizing shadows that could reveal the insect's outline. This principle is observed in stick insects and grasshoppers, where it counters the overhead light typical of open habitats. Chemical defenses provide insects with potent means to repel or harm attackers through the production or sequestration of toxins. Monarch butterflies (Danaus plexippus) exemplify this by sequestering cardenolides from milkweed plants during their larval stage, rendering adults unpalatable or toxic to vertebrates like birds. This sequestration not only deters predation but also persists through metamorphosis, with empirical tests showing reduced attack rates on dosed models. Other insects, such as bombardier beetles, synthesize their own irritants, ejecting hot quinone sprays from abdominal glands to startle predators. Behavioral defenses enable rapid responses to immediate threats without relying on morphology or chemistry. Autotomy, the voluntary detachment of appendages like legs, is common in spiders and insects such as crickets, allowing escape from grasping predators while the discarded limb distracts the attacker. Regrowth occurs in subsequent molts, though at an energetic cost. Thanatosis, or feigning death, involves insects like certain ground beetles assuming a rigid, immobile posture to mimic carrion, exploiting predators' aversion to spoiled prey; this tactic is triggered by tactile or vibrational cues and can last minutes to hours. Aposematism uses conspicuous warning signals to advertise defenses, promoting predator learning and avoidance. Ladybugs (Coccinellidae) display bold red-and-black patterns paired with alkaloid-based unpalatability, where bitter hemolymph deters feeding; field experiments confirm that predators like birds quickly associate these colors with distastefulness after initial encounters. This strategy often evolves alongside mimicry, where harmless species copy the signals of defended models to gain protection. Chemical signals, such as alarm pheromones, may reinforce aposematism by alerting conspecifics to dangers.

Human Interactions

Agricultural and medical impacts

Insects exert profound negative influences on global agriculture through direct crop damage, post-harvest losses, and ecosystem disruptions caused by invasive species. Among the most devastating are locust swarms, which can rapidly consume vast areas of vegetation; the 2020 East African desert locust upsurge, the worst in decades, affected over 21 million people by destroying crops and pastures across Ethiopia, Kenya, Somalia, and other nations, leading to vegetation and crop losses of 42% to 69% in vulnerable regions.^[107]^[108] Similarly, the boll weevil (Anthonomus grandis) has historically ravaged cotton production in the southeastern United States; following its arrival in 1892, it caused cotton yields to plummet by up to 50% within five years of infestation in affected counties, reducing total acreage from 5.2 million to 2.6 million acres between 1914 and 1923.^[109]^[110] Stored product pests, such as grain weevils (Sitophilus spp.), further compound agricultural losses by infesting harvested commodities during storage, resulting in quality degradation and quantity reductions. These insects are responsible for an estimated 10% of global grain production losses annually, with developing countries experiencing up to one-third of their stored grain destroyed each year due to such infestations.^[111]^[112] The economic toll is substantial, contributing to billions in global losses through direct damage, contamination, and the costs of control measures.^[113] Invasive insect species amplify these agricultural threats by targeting native flora and forestry resources. The emerald ash borer (Agrilus planipennis), introduced from Asia, has killed hundreds of millions of ash trees across North America since its detection in 2002, with larval feeding girdling phloem tissue and causing mortality in as little as two years for infested trees.^[114]^[115] This devastation threatens urban and rural landscapes, with potential losses to the U.S. timber industry alone exceeding $10 billion from the estimated 7.5 billion ash trees at risk.^[116] Beyond agriculture, insects pose significant medical risks as vectors for debilitating diseases, transmitting pathogens that affect human health on a massive scale. Mosquitoes, particularly Anopheles species, are primary vectors for malaria, caused by Plasmodium parasites; in 2023, this disease resulted in 263 million cases and 597,000 deaths worldwide, predominantly in sub-Saharan Africa.^[117]^[118] Ticks, such as the blacklegged tick (Ixodes scapularis), transmit Lyme disease via Borrelia burgdorferi bacteria during prolonged feeding, with transmission typically requiring 24-48 hours of attachment; in the United States, this leads to over 476,000 estimated cases annually, causing symptoms ranging from fever and rash to severe neurological complications if untreated.^[119]^[120] These vector-borne illnesses not only strain healthcare systems but also incur substantial economic burdens through treatment, lost productivity, and preventive efforts.^[121]

Beneficial uses and conservation

Insects provide essential services to human agriculture and ecosystems, most notably through pollination, which supports the production of fruits, vegetables, and nuts. Globally, insect pollinators, particularly bees, contribute an estimated $235–$577 billion annually (as of 2025) to crop values by facilitating the reproduction of approximately 75% of leading food crops.^[122]^[123] This economic valuation underscores the irreplaceable role of insects in sustaining food security, as their decline could disrupt yields of key commodities like coffee, cocoa, and almonds.^[124] Another critical beneficial use involves biological control, where predatory or parasitic insects are deployed to manage pest populations without synthetic chemicals. Trichogramma wasps, tiny egg parasitoids, exemplify this approach by targeting the eggs of lepidopteran pests such as moths and butterflies that damage crops like corn, cotton, and sugarcane.^[125] These wasps have been mass-reared and released in biological control programs worldwide, including in China and the United States, reducing pest infestations by up to 90% in some field trials while minimizing environmental harm from pesticides.^[126] Despite these benefits, insect populations face severe conservation challenges, with over 40% of species threatened by extinction due primarily to habitat loss from urbanization and agriculture, as well as pesticide exposure.^[127] As of 2025, studies continue to report that over 40% of insect species face extinction risks due to these factors.^[128] Neonicotinoid insecticides, widely used in farming, exacerbate these declines by impairing insect navigation, reproduction, and immunity, leading to reduced abundances of bees, butterflies, and aquatic insects even at sublethal doses.^[129] The 2019 IPBES Global Assessment highlighted that around 1 million species overall, including a significant portion of insects, are at risk of extinction within decades, driven by these anthropogenic pressures. Conservation initiatives aim to mitigate these threats through habitat protection and restoration. For instance, the Monarch Butterfly Biosphere Reserve in Mexico, a UNESCO World Heritage site spanning over 56,000 hectares, safeguards critical overwintering grounds for monarch butterflies (Danaus plexippus), supporting their migration and breeding while promoting sustainable ecotourism. In North America, programs like those from the U.S. Fish and Wildlife Service and Natural Resources Conservation Service provide incentives for landowners to create milkweed-rich habitats, helping counter the 80-90% decline in eastern monarch populations since the 1990s.^[130] These efforts emphasize integrated strategies, such as reducing pesticide use and enhancing connectivity between protected areas, to bolster insect resilience amid ongoing environmental changes.

Cultural and scientific significance

Insects have held profound cultural significance across civilizations, often symbolizing transformation, rebirth, and the divine. In ancient Egyptian mythology, the scarab beetle (Scarabaeus sacer) represented the sun god Khepri, embodying the cycle of renewal as the beetle was observed rolling dung balls, mirroring the sun's daily journey across the sky.^[131] Butterflies, with their metamorphic life cycle, have appeared in global folklore as emblems of the soul and spiritual change; in ancient Greek tradition, the term "psyche" denoted both the human soul and the butterfly, linking the insect to immortality and the afterlife.^[132] Humans have long derived valuable products from insects, integrating them into economies and industries. Silkworms (Bombyx mori) produce silk, a fiber harvested from cocoons that has been central to textile production for millennia, originating in ancient China around 3500 BCE.^[133] Honeybees (Apis mellifera) yield honey and beeswax, used since prehistoric times for food, medicine, and candles, with honey serving as a natural sweetener and antimicrobial agent.^[133] Cochineal insects (Dactylopius coccus) provide carminic acid, extracted to create the vibrant red dye carmine, which colored textiles, cosmetics, and foods in Mesoamerican cultures and later globally.^[133] Lac bugs (Kerria lacca) secrete resin processed into shellac, a versatile coating for wood finishes, pharmaceuticals, and varnishes, dating back to ancient India.^[133] Over 2,200 insect species are consumed worldwide as food, offering a sustainable protein source amid growing global demand.^[134] Crickets (Acheta domesticus), for instance, contain approximately 60-70% protein by dry weight, surpassing many traditional meats in amino acid completeness and providing essential micronutrients like iron and B vitamins.^[135] In scientific research, insects serve as pivotal model organisms; the fruit fly Drosophila melanogaster has been instrumental in genetics since Thomas Hunt Morgan's 1910 experiments, enabling discoveries in inheritance, mutation, and developmental biology that earned Nobel Prizes.^[136] Recent advances in genetic engineering, such as CRISPR-Cas9 applications in mosquitoes (Anopheles gambiae), have progressed in 2025 to develop self-limiting gene drives that induce female sterility to curb malaria transmission without permanent ecological alteration.^[137] Biomimicry draws from insect behaviors for innovation, with ant colony optimization algorithms— inspired by pheromone trails in species like Argentine ants (Linepithema humile)—applied in robotics for efficient path planning and swarm coordination since their introduction in 1992.^[138]

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An open system, like the insects, has problems with circulation. To aid in hemolymph movement, insects have evolved a dorsal and ventral diaphragm, which ...
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The mosquito circulatory system
The primary organ driving hemolymph circulation in mosquitoes is a dorsal vessel that is subdivided into an abdominal heart and a thoracic aorta.
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Respiratory System – ENT 425 – General Entomology
Small insects rely almost exclusively on passive diffusion and physical activity for the movement of gasses within the tracheal system. However, larger insects ...
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5.1 Systems of Gas Exchange – Animal Physiology
Air enters and leaves the tracheal system through the spiracles. Some insects can ventilate the tracheal system with body movements. Figure 5.6. Insects ...
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Animal Gas Exchange and Transport | Organismal Biology
Air enters and leaves the tracheal system through the spiracles. Some insects can ventilate the tracheal system with body movements. Insects perform respiration ...
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Reproductive System – ENT 425 – General Entomology
The reproductive organs of insects are similar in structure and function to those of vertebrates: a male's testes produce sperm and a female's ovaries produce ...
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[PDF] Ecology and physiology - Information Technology Solutions
Juvenile hormone is a major hormone for this adult insect. JH is essential ... Accessory reproductive glands and male reproductive tract. 2. High titers ...
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[PDF] Unit 4 in Entomology
Two of the molting hormones we'll discuss are juvenile hormone, also called J. H., and ecdysone. Juvenile hormone is secreted from a part of an insect brain ...
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Mating in Insects | Smithsonian Institution
Insect mating involves locating mates via pheromones, courtship rituals, and internal fertilization, with some exceptions like bedbugs.
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Insect Mating Behaviors: A Review of the Regulatory Role of ...
May 8, 2025 · In our findings, we note that the mating process of insects entails mate attraction, courtship, copulation, and post-mating behaviors. The 18 ...
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Honeybee drones are attracted by groups of consexuals in a ...
When a receptive female, a queen, enters the congregation, drones are attracted to her by queen-produced pheromones and visual cues and attempt to mate with the ...
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Flash signal evolution, mate choice, and predation in fireflies
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Aggression and Courtship in Drosophila: Pheromonal ...
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SPSL1 is essential for spermatophore formation and sperm ... - NIH
Dec 4, 2023 · In lepidopteran females, the spermatophore consists of proteins from the seminal fluid that are transferred from males during copulation [23].
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Traumatic insemination and sexual conflict in the bed bug Cimex ...
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Costly traumatic insemination and a female counter-adaptation ... - NIH
Male bed bugs pierce females through the body wall and inseminate directly into the body cavity. It has previously been shown that such traumatic insemination ...
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Genetic variation in paternal investment in a seed beetle
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Biparental care in insects: Paternal care, life history, and the function ...
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Sexual Selection and Paternal Investment in Insects
Trivers (1972) defines parental investment as "any investment by the parent in an individual offspring that increases the offspring's chance of surviving (and ...
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The evolution of insect metamorphosis: a developmental and ...
Aug 26, 2019 · In contrast to holometabolous species, the embryos of ametabolous and hemimetabolous species are quite sensitive to treatment with ...
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Juvenile hormone regulates body size and perturbs insulin signaling ...
Apr 28, 2014 · In insects, the hormones ecdysone and juvenile hormone (JH) control the timing of the metamorphic transition and hence growth duration (1).
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An Introduction to Eusociality | Learn Science at Scitable - Nature
The order Hymenoptera is the largest and most well-known animal group with eusocial species (see Figures 1–3). Most hymenoptera are not eusocial, but this ...
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[PDF] Evolution of eusociality and the soldier caste in termites
This caste is ancestral in termites and evolved prior to a true worker caste (i.e. a sterile helping caste). Unlike the soldiers found in the other eusocial ...
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Hamilton's rule and the causes of social evolution - PubMed Central
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Division of labor in honeybees: form, function, and proximate ...
Honeybees exhibit two patterns of organization of work. In the spring and summer, division of labor is used to maximize growth rate and resource accumulation.
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ENY-166/IN1102: The Social Organization of Honey Bees
Reproductive division of labor—Each honey bee colony consists of a queen (reproductive female), workers (non-reproductive females), and drones (males). Each ...Missing: hives | Show results with:hives
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Review Pheromone production in bark beetles - ScienceDirect.com
Bark beetles use aggregation pheromones to coordinate mass attacks on host pine. Feeding induces production of an aggregation pheromone in the pioneer sex ...
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[PDF] Aggregation in non-social insects
Several bark beetle species also show dose-dependent response to aggregation pheromones released by conspecifics (reviewed in Raffa, 2001). Fig. 1. Percent ...
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Meeting the challenges of on-host and off-host water balance ... - NIH
Typically, water in tissues is conserved at the expense of the hemolymph, as exemplified by the tenebrionid beetle, Onymacris plana, a species that loses ...
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Respiration in Aquatic Insects - ENT 425 - NC State University
This rectal gill mechanism doubles as a jet propulsion system. A sudden, powerful contraction of the abdomen will expel a jet of water and thrust the insect ...<|separator|>
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How do insects survive the winter? - Integrated Crop Management
Mar 4, 2016 · Some freeze-avoidant species acclimate for winter by creating cryoprotectants, or antifreeze proteins, in their body (usually 20-30% of ...
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Cockroaches Management Guidelines - UC IPM
Most cockroaches harbor within moist, dark crevices when not foraging for food. They crawl quickly and may climb rough surfaces.
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The Importance of Pollinators - USDA
Pollinators by Numbers. Three-fourths of the world's flowering plants and about 35 percent of the world's food crops depend on animal pollinators to reproduce.
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[PDF] How many flowering plants are pollinated by animals?
Some of the most cited sources, whose estimates range from 67% to 96% of all angiosperm species being animal pollinated, are obscure as to how these figures ...
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The importance of insects on land and in water: a tropical view
### Summary of Insects in Nutrient Cycling and Decomposition
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The Ecology of Carrion Decomposition | Learn Science at Scitable
Decomposition of animal carrion is achieved primarily through the activities of invertebrates, such as flies and beetles, and large scavengers.
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Identification of Conditions for Successful Aphid Control by ... - NIH
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Uncovering the Potential of Termite Gut Microbiome for ... - Frontiers
This study demonstrates that termite-gut microbiomes constitute a reservoir of lignocellulose-degrading bacteria that can be harnessed in artificial conditions.
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Starting in late 2019, the largest upsurge of desert locusts in living memory darkened the skies of East Africa and Yemen, threatening to wipe out crops...
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An Integrated Assessment Approach for Socio‐Economic ...
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The Impact of the Boll Weevil, 1892–1932 | The Journal of Economic ...
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Boll Weevil - New Georgia Encyclopedia
Yield losses associated with the boll weevil reduced cotton acreage from a historical high of 5.2 million acres during 1914 to 2.6 million acres in 1923.
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Tackling insect infestation and quality loss: A review - ScienceDirect
Research indicates that insect-related damage during storage can result in economic losses of up to 10 % of global grain production, with developing nations ...
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The Real Costs of Stored Product Pests
Dec 8, 2017 · An estimated 1/4 to 1/3 of the world grain crop is lost each year in storage. · Stored product pests are responsible for millions of dollars of ...
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The Effects of Insect Infestation on Stored Agricultural Products and ...
May 18, 2023 · In this review article, we focus on the effects of insect pests on the quality of stored cereals and legume grains.
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Emerald Ash Borer | National Invasive Species Information Center
The emerald ash borer, an invasive beetle native to Asia, is one of the most destructive invasive species in North America.
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Impacts of the emerald ash borer (Agrilus planipennis Fairmaire ...
The larval feeding behavior of the emerald ash borer (EAB) effectively girdles the tree's phloem tissue resulting in tree mortality in as little as 2 years.
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As the emerald ash borer spreads, the damage will mount: an estimated 7.5 billion ash trees are found on timberlands in US (USDA APHIS 2007); an additional 30- ...
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Apr 2, 2025 · An estimated 263 million cases of malaria and 597,000 deaths from malaria occurred worldwide in 2023. Nearly half the global population lives in ...
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How Lyme Disease Spreads - CDC
Sep 24, 2024 · Lyme disease bacteria causing human infection in the United States are spread to people by blacklegged (Ixodes) ticks.
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Lyme Disease - CDC
May 14, 2024 · Lyme disease is caused by Borrelia bacteria spread to people by the bite of an infected blacklegged tick.
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Analysis Economic valuation of the vulnerability of world agriculture ...
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Economic Value Of Insect Pollination Worldwide Estimated At U.S. ...
Sep 15, 2008 · The worldwide economic value of the pollination service provided by insect pollinators, bees mainly, was €153 billion in 2005 for the main crops that feed the ...
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Trichogramma Wasp | NC State Extension - Entomology
Trichogramma wasps can contribute significantly to natural control of a variety of insect populations. They are probably the most widely released insect for ...
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Jan 7, 2021 · Trichogramma species make up one of the most commonly used groups of natural enemies for biological control programs worldwide.
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Worldwide decline of the entomofauna: A review of its drivers
Over 40% of insect species are threatened with extinction. •. Lepidoptera, Hymenoptera and dung beetles (Coleoptera) are the taxa most affected. •. Four ...
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Experimental evidence for neonicotinoid driven decline in aquatic ...
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Here, you can find information about how you can help monarchs, contribute to their habitat and find resources and assistance to help guide your actions.
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McClung Museum - Egyptian Scarabs
The scarab-beetle was the symbol of the Sun-god and as such could stimulate the deceased's heart to life. · The scarab-beetle was the symbol of "transformations, ...
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Folklore of Butterflies - SeasonWords.com
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From Silk Moths to Fruit Flies, These Five Insects Have Changed the ...
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The global atlas of edible insects: analysis of diversity and ... - Nature
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From Farm to Fork: Crickets as Alternative Source of Protein ...
It has been postulated that crickets generally contain high-quality nutrients, which are easily digestible and more bio-available than those available from ...
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Jun 4, 2015 · From its first use in the laboratory in the early 1900s until the present day, Drosophila melanogaster has been central to major breakthroughs in genetics.
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Aug 7, 2025 · The Ant Colony Optimization (ACO) is a population-based metaheuristic algorithm for the optimization problem, inspired by the foraging behavior ...