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Polyphaga

Polyphaga is the largest and most diverse suborder within the insect order Coleoptera, commonly known as beetles, comprising approximately 90% of all beetle species—over 350,000 described species across approximately 150 families organized into 16 superfamilies (total Coleoptera ~400,000 species).^[1] This suborder is distinguished morphologically by its hind coxae, which do not divide the first abdominal segment, in contrast to the suborder Adephaga.^[2] Polyphagans exhibit extraordinary adaptability, occupying diverse habitats from freshwater and soil to plants, dung, and fungi, with body sizes ranging from less than 1 mm to over 100 mm.^[1] Their ecological roles span predation, herbivory, decomposition, and pollination, making them integral to ecosystems worldwide.^[2] Taxonomically, Polyphaga is divided into several series and infraorders, including Staphyliniformia (rove beetles and allies), Elateriformia (click beetles and relatives), Scarabaeiformia (scarab beetles), Bostrichiformia (deathwatch and skin beetles), and Cucujiformia (lady beetles, longhorn beetles, and weevils), reflecting a broad spectrum of evolutionary divergence.^[1] The suborder has a robust fossil record dating back to the Early Triassic, with significant diversification evident from the Jurassic onward, particularly in phytophagous lineages.^[1]^[3] Dominant families such as Curculionidae (weevils, ~50,000 species) and Scarabaeidae (scarabs, ~25,000 species) highlight its species richness, with many groups specialized as wood-borers or leaf-feeders.^[1] Polyphaga's diversity underscores its evolutionary success, representing about 30–40% of all described insect species and influencing agriculture, forestry, and biodiversity conservation.^[4] Notable examples include beneficial predators like ladybird beetles (Coccinellidae), which control aphids, and economically damaging pests such as the boll weevil (Anthonomus grandis) in the family Curculionidae.^[2] Larval forms vary widely, from campodeiform (active, legged) to eruciform (grublike, legless), adapting to specific niches like soil-dwelling or plant-mining.^[2] Ongoing research emphasizes Polyphaga's role in understanding insect evolution, with genomic studies revealing adaptations for plant-feeding diets in many lineages.^[5]

Taxonomy and systematics

Etymology

The name Polyphaga derives from the Ancient Greek roots poly- (πολύς, meaning "many") and phagein (φαγεῖν, meaning "to eat"), collectively signifying "many-eaters" or "eaters of many things," in reference to the suborder's exceptionally broad range of feeding strategies that incorporate plant, animal, and fungal resources.^[6] This etymological choice underscores the group's dietary versatility relative to the more specialized, predominantly carnivorous and aquatic feeding observed in the suborder Adephaga.^[2] The suborder was formally established by Italian entomologist Carlo Emery in 1886 as part of his classification of Coleoptera.^[7]

Classification

Polyphaga is one of four suborders within the order Coleoptera, the others being Archostemata, Adephaga, and Myxophaga. It represents the most diverse suborder, encompassing the vast majority of beetle species and exhibiting extensive morphological and ecological variation.^[8] The suborder is divided into five series: Bostrichiformia, Staphyliniformia, Scarabaeiformia, Elateriformia, and Cucujiformia. These series collectively include 16 superfamilies and approximately 144 families. Notable superfamilies encompass Scarabaeoidea (scarab and stag beetles), Curculionoidea (weevils), Tenebrionoidea (darkling beetles and allies), Staphylinoidea (rove beetles), and Elateroidea (click and fireflies beetles), among others. Approximate species richness varies by group, with Curculionoidea comprising around 62,000 species, Chrysomeloidea (leaf and longhorn beetles) about 50,000 species, and Staphylinoidea over 48,000 species; overall, Polyphaga contains roughly 350,000 described species.^[7]^[8]^[3] Classification at the subordinal level relies on key diagnostic traits in adults, including a tarsal formula of 5-5-5 (five tarsomeres on each tarsus) and the absence of a notopleural suture on the prothorax. These features contrast with other suborders, such as Adephaga, which typically exhibit a 5-5-4 or 4-4-4 tarsal formula and a visible notopleural suture.^[9]^[10]

Phylogenetic position

Polyphaga constitutes the largest suborder of Coleoptera, comprising over 90% of all beetle species, and occupies a basal position in the order's phylogeny as the sister group to the clade comprising Adephaga, Myxophaga, and Archostemata, with Adephaga sister to the [Myxophaga + Archostemata] clade. This configuration is robustly supported by molecular phylogenies integrating transcriptomic and genomic data, including large-scale studies employing hundreds of nuclear loci across diverse beetle taxa.^[11]^[8] Key synapomorphies defining Polyphaga include the loss of the basal portion of the radial sector (Rs) vein in the hindwings, which distinguishes it from more basal suborders, and the development of flexible intersegmental connections in the abdomen that enhance mobility. These morphological traits are evident in both adult and larval stages and have been corroborated through comparative anatomical studies of wing venation and abdominal sclerites.^[12]^[13] Within Polyphaga, the monophyly of major series such as Elateriformia, Cucujiformia, and Bostrichiformia is generally well-supported, but debates persist regarding the integrity of Staphyliniformia, which recent genomic datasets indicate is paraphyletic. Specifically, Scarabaeiformia (including scarab beetles) nests within traditional Staphyliniformia as sister to Staphylinoidea, necessitating taxonomic revisions to restore monophyly, as proposed in integrated phylogenomic frameworks. These findings highlight ongoing challenges in resolving deep divergences among polyphagan series using both molecular and morphological evidence.^[11]

Evolutionary history

Origins

The suborder Polyphaga, comprising the largest and most diverse group within the order Coleoptera, is believed to have originated in the Late Permian period, approximately 260 million years ago, based on rare fossil evidence and phylogenetic analyses. The earliest records consist of fragmentary remains and trace fossils, such as wood-borings attributed to polyphagan-like xylophagous beetles, indicating their presence in late Paleozoic ecosystems dominated by gymnosperm forests.^[14] These primitive forms exhibit morphological features, including elongated bodies and elytral structures, that resemble those of modern Elateriformia, the basal series within Polyphaga, suggesting an early diversification among wood-associated lineages.^[15] A significant radiation of Polyphaga occurred during the Triassic-Jurassic transition, roughly 200 to 150 million years ago, marking the shift from rare, stem-group dominated assemblages to more diverse crown-group forms. This period saw the emergence of key superfamilies within series such as Elateriformia and Staphyliniformia, as evidenced by fossils from lacustrine and amber deposits.^[15] The timing coincides with the initial evolution of angiosperms in the Late Jurassic and the ongoing breakup of the supercontinent Pangaea, which began fragmenting in the Late Triassic and facilitated biogeographic dispersal and ecological opportunities in recovering terrestrial habitats.^[11] The persistence of Polyphaga through major geological events, including the end-Permian mass extinction—the most severe in Earth's history—was facilitated by relatively low extinction rates compared to other insect groups. Genomic studies indicate that early polyphagans, adapted to detrital and wood-based niches, experienced minimal lineage turnover during this crisis, allowing survival and subsequent diversification as ecosystems rebounded in the Mesozoic.^[16] This resilience is supported by molecular clock estimates placing the divergence of Polyphaga within Coleoptera around the Carboniferous-Permian boundary, with crown-group radiation postdating the extinction event.^[17]

Diversification and fossil record

The diversification of Polyphaga began to accelerate following their Permian origins, with early evidence appearing in the Triassic. Fossil impressions suggestive of polyphagans have been identified in the Madygen Formation of Kyrgyzstan, dating to the Middle-Late Triassic (approximately 242–228 Ma), including members of the extinct family Peltosynidae and undescribed elaterids that indicate basal polyphagan morphologies adapted to lacustrine environments.^[18]^[19] These rare Triassic records highlight a sparse but foundational presence, contrasting with the dominance of archostematan beetles at the time.^[15] A major radiation occurred during the Jurassic (approximately 201–145 Ma), marked by the explosive diversification of Elateriformia and Cucujiformia, which together represent key basal lineages within Polyphaga. Elateriformia fossils from sites like the Daohugou Beds in China include early dascilloids, buprestids, and elateroids, such as Artematopodidae and Cerophytidae, demonstrating series-level diversity in click and metallic wood-boring beetles.^[15] Similarly, Cucujiformia saw the emergence of cleroideans (e.g., Melyridae, Cleridae) and early cucujoids, with well-preserved specimens in Jurassic compressions from Karatau in Kazakhstan revealing adaptations for predatory and fungivorous lifestyles.^[15] Jurassic compression fossils, such as those from the Upper Jurassic Talbragar Fish Bed in Australia, further document elateroid diversity, including the basal genus Wongaroo, underscoring the suborder's expansion across Gondwanan and Laurasian landmasses. The Cretaceous (approximately 145–66 Ma) witnessed a pronounced boom in herbivorous polyphagan groups, particularly within Scarabaeoidea, closely tied to the radiation of flowering plants (angiosperms). Scarab fossils from Lower Cretaceous deposits, such as the Yixian Formation in China (e.g., Septiventer quadridentatus), show early diversification of phytophagous and coprophagous lineages, with crown-group pleurostict scarabs and Glaphyridae originating between 108–141 Ma, coinciding with angiosperm proliferation around 132–141 Ma.^[20] This period's amber deposits, including Burmese and Lebanese ambers, preserve diverse polyphagans like tenebrionoids and curculionoids, illustrating ecological shifts toward angiosperm-dependent feeding. Overall, net diversification rates remained positive and low throughout Polyphaga's history, with family-level extinction rates at zero, even across the Cretaceous–Paleogene boundary.^[21] By the Eocene (approximately 56–33 Ma), over 90% of modern Polyphaga diversity at the family level had been achieved, as evidenced by rich amber and compression fossils from sites like the Baltic and Green River formations, which capture nearly all extant superfamilies with minimal subsequent turnover.^[21] This stabilization reflects sustained speciation driven by ecological opportunism, with 63% of living families already represented in the fossil record by this epoch.^[21]

Morphology

Adult features

Adult Polyphaga beetles are characterized by their hardened forewings, known as elytra, which cover and protect the folded membranous hindwings beneath them. These elytra exhibit variations in fusion along the midline suture; for instance, in weevils (superfamily Curculionoidea), the elytra are often completely fused, forming a seamless shield, while in click beetles (family Elateridae), they remain loosely articulated, allowing rapid opening for the beetle's characteristic jumping mechanism.^[22] Additionally, the prothorax in Polyphaga lacks a distinct notopleural suture, a feature that distinguishes this suborder from Adephaga, where such a suture is present on the underside of the pronotum. The antennae of adult Polyphaga display considerable diversity in form, ranging from filiform (thread-like) in many ground-dwelling species to lamellate (plate-like) in scarab beetles (family Scarabaeidae), aiding in sensory functions such as chemoreception. Mouthparts are primarily adapted for chewing, with robust mandibles typical across the suborder, though specialized modifications occur; notably, in the superfamily Curculionoidea, the head is prolonged into an elongated rostrum at the apex of which the chewing mouthparts are located, facilitating precise feeding on plant tissues.^[23]^[2]^[24] The tarsal formula in adult Polyphaga is typically 5-5-5, though reduced in some groups (e.g., 5-5-4 or 4-4-4), meaning each leg usually has five tarsal segments, a trait that aids in identification and locomotion across diverse habitats. Sexual dimorphism in tarsal structure is evident in some families, such as Scarabaeidae, where males often possess inflated or broadened fore tarsi adapted for grasping during mating.^[23]^[25]

Larval characteristics

The larvae of Polyphaga display considerable morphological variability adapted to diverse microhabitats and feeding strategies, ranging from C-shaped eruciform forms in wood-boring species to elongate, active campodeiform forms in predatory ones. Eruciform larvae, characterized by a fleshy, curved body with reduced sclerotization and thoracic legs often absent or vestigial, are typical of families like Buprestidae, where they facilitate boring into wood or plant tissues.^[26] In contrast, campodeiform larvae, with a flattened, sclerotized body, prominent legs, and elongated cerci, predominate in predatory groups such as Staphylinidae, enabling rapid movement and prey capture.^[27] The head capsule is prognathous and well-sclerotized across Polyphaga larvae, featuring a variable number of stemmata—typically 0 to 6 per side—depending on the family and ecological niche, which aids in navigation and prey detection.^[28] Thoracic legs are generally present and functional in active larval forms like campodeiform types, consisting of three pairs with one- or two-clawed tarsi, but they are reduced or absent in more sedentary eruciform or scarabaeiform larvae specialized for burrowing or feeding in concealed spaces. Abdominal segmentation is distinct, with 10 segments visible, though the ninth often bears specialized structures such as urogomphi—paired, horn-like projections surrounding a caudal notch—in families like Elateridae, where wireworm larvae use them for anchorage and locomotion in soil.^[29] These urogomphi vary in length and shape, contributing to the larvae's ability to navigate tight tunnels. Specific adaptations reflect larval lifestyles, including hardened terga (dorsal sclerites) in soil-dwelling forms like elaterid wireworms, providing protection against desiccation and predation in subterranean environments.^[30] In species with fecal shields, such as certain Chrysomelidae (e.g., Cassidinae), larvae construct protective structures from their excrement, held over the body via an anal sling to shield against predators and parasites during vulnerable feeding stages. These features underscore the suborder's evolutionary flexibility in larval morphology.

Biology and ecology

Life cycle

Polyphaga, like all beetles, undergo holometabolous (complete) metamorphosis, consisting of four distinct life stages: egg, larva, pupa, and adult. The egg stage typically lasts from a few days to several weeks, depending on species and environmental conditions, with females laying clusters of eggs in protected locations. Larvae, often referred to as grubs or worms, pass through multiple instars—ranging from 3 to more than 20—during which they grow rapidly through molting; this stage can endure from weeks to years, varying widely among families. The pupal stage occurs within a cocoon, earthen cell, or chamber formed by the final instar larva, lasting days to months as the insect undergoes reorganization into the adult form. Adults emerge through ecdysis, ready for reproduction, though some temperate species enter diapause during larval or pupal stages to overwinter, synchronizing development with favorable seasons.^[31]^[32]^[32] Reproduction in Polyphaga involves internal fertilization, where males transfer sperm via aedeagus during copulation, followed by females ovipositing fertilized eggs in substrate-specific sites suited to larval needs. For instance, in Scarabaeidae (scarab beetles), eggs are commonly deposited in soil, often near plant roots or decaying matter, with oviposition influenced by soil moisture and texture. In contrast, Cerambycidae (longhorn beetles) females chew slits in bark or wood to lay eggs individually, ensuring larvae have access to woody tissues for development. These strategies reflect adaptations to diverse ecological niches, with egg numbers varying from dozens to hundreds per female across species.^[33]^[34]^[35] Parental care is rare in Polyphaga but occurs in select taxa, such as certain Staphylinidae (rove beetles), where subsocial behaviors include egg guarding and provisioning for early-instar larvae in burrows or chambers. For example, in Bledius spectabilis, females protect eggs and young larvae from predators in intertidal mud burrows, enhancing offspring survival. Generation times span from weeks in fast-developing predatory species, like some Coccinellidae (lady beetles) that complete cycles in 3–4 weeks under optimal conditions, to several years in wood-boring groups such as Cerambycidae, where larval development alone can take 1–3 years depending on host quality and climate. These variations underscore the suborder's adaptability to temporal environmental cues.^[36]^[37]^[38]

Habitat and feeding habits

Polyphaga beetles occupy a wide array of terrestrial microhabitats, including soil, leaf litter, wood, and vegetation, with some species specialized for aquatic environments such as ponds, streams, and wetlands. While predominantly terrestrial, polyphagan families like Hydrophilidae and Elmidae inhabit standing waters and flowing streams, where they exploit algae-covered substrates or submerged vegetation.^[39]^[40] The suborder exhibits remarkable dietary diversity, encompassing multiple feeding guilds that reflect adaptations to various trophic levels. Herbivorous species, such as leaf beetles (Chrysomelidae), primarily consume foliage and plant tissues, contributing to herbivory across forests and grasslands. Detritivores like dung beetles (Scarabaeidae) break down animal feces, recycling nutrients in ecosystems. Predatory polyphagans, including rove beetles (Staphylinidae), hunt small invertebrates in soil and litter, regulating pest populations. Fungivorous bark beetles (Curculionidae: Scolytinae) feed on fungal mycelium within tree phloem, often in decaying or stressed wood.^[16]^[9]^[39] Behavioral adaptations in Polyphaga enhance survival in these niches, particularly through defense mechanisms. Blister beetles (Meloidae) secrete cantharidin, a toxic blistering agent from their hemolymph, deterring predators upon disturbance. Mimicry is evident in longhorn beetles (Cerambycidae), such as Clytus arietis, which adopt black-and-yellow coloration and jerky movements to resemble stinging wasps, exploiting Batesian mimicry for protection.^[41]^[42] Microhabitat specialization often involves symbiotic relationships, as seen in ambrosia beetles (Scolytinae), which excavate galleries in wood and cultivate mutualistic fungi as their primary food source, with the fungi detoxifying the substrate and providing nutrients. These interactions enable exploitation of nutrient-poor environments like xylem. Morphological features, such as robust mandibles, support these feeding strategies across guilds.^[43]^[44]

Diversity and distribution

Species richness

Polyphaga is the most species-rich suborder of Coleoptera, encompassing approximately 360,000 described species as of 2025, which accounts for 90% of all known beetle diversity.^[45] Estimates suggest the true total, including undescribed species, may reach 1 to 2 million, reflecting the suborder's vast untapped diversity driven by ongoing taxonomic efforts and molecular discoveries.^[46] This immense richness is unevenly distributed across its taxonomic hierarchy, with a few families dominating the count. The series Cucujiformia contributes the largest share, comprising around 60% of Polyphaga species, primarily through phytophagous groups like weevils and leaf beetles.^[47] Staphyliniformia follows with approximately 20%, largely due to the hyperdiverse rove beetles (Staphylinidae) alone, which include over 66,000 described species and potentially far more undescribed ones, especially in tropical regions.^[48]^[49] Other series, such as Elateriformia and Scarabaeiformia, make up the remainder, with Scarabaeidae contributing about 36,000 species focused on dung-feeding and phytophagous habits.^[50] Among the most species-rich families are Curculionidae (weevils) with roughly 62,000 species, Chrysomelidae (leaf beetles) with about 37,000 species, and Staphylinidae as noted.^[51]^[52] Undescribed diversity is particularly pronounced in tropical weevils and rove beetles, where habitat complexity and morphological conservatism hinder traditional taxonomy.^[53] Recent trends in species discovery highlight the role of genomic approaches in uncovering hidden diversity within Polyphaga. For instance, phylogenomic surveys in the 2020s have revealed cryptic species complexes in Tenebrionidae (darkling beetles), such as in Atacama Desert lineages, emphasizing underestimated speciation driven by environmental adaptation.^[54] These findings underscore how molecular tools are accelerating the documentation of Polyphaga's richness, potentially elevating family-level counts in the coming years.

Global patterns

Polyphaga, the largest suborder of beetles, exhibits a cosmopolitan distribution, occurring on all continents except Antarctica and the extreme polar regions of the Arctic, where harsh conditions limit their presence.^[55] This suborder thrives in diverse environments, from forests and grasslands to deserts and freshwater systems, reflecting its adaptive versatility across terrestrial and semi-aquatic habitats. Diversity gradients follow latitudinal patterns typical of insects, with species richness peaking in tropical regions and declining toward temperate and arid zones.^[56] The tropics harbor the highest concentrations of Polyphaga species, driven by historical radiations linked to angiosperm diversification and stable climates. In the Neotropics, particularly the Amazon basin, this suborder contributes substantially to global beetle diversity through extensive herbivorous and detritivorous lineages. Endemism is pronounced in isolated hotspots, such as Madagascar, where nearly all dung beetle species (Scarabaeinae, a Polyphaga superfamily) are endemic, representing over 96% of the ~300 described taxa resulting from multiple independent colonizations and radiations. Similarly, Australia hosts high levels of endemism in buprestid beetles (Buprestidae), with over half of its genera and most species restricted to the continent, showcasing biogeographic peculiarities tied to its unique flora and isolation.^[56]^[57]^[58] Biogeographic patterns in Polyphaga are shaped by both ancient vicariance and more recent dispersal mechanisms. Gondwanan vicariance has influenced southern hemisphere families, such as certain water scavenger beetles (Hydrophilidae), where mid-Cretaceous fragmentation of West Gondwana (Africa-South America split ~100 Ma) drove early diversification and disjunct distributions. Dispersal occurs via wind currents, rafting on vegetation, and human activities, enabling range expansions beyond natural barriers. A prominent example is the khapra beetle (Trogoderma granarium, Dermestidae), a cosmopolitan stored-product pest native to the Indian subcontinent and Middle East, which has invaded over 60 countries across Africa, Asia, Europe, and North America through global trade in grains and commodities, establishing populations where environmental suitability allows.^[59]^[60]^[61]

Human interactions

Economic roles

Polyphaga beetles exert significant negative economic impacts through their role as pests in agriculture and forestry. Bark beetles in the subfamily Scolytinae, such as the mountain pine beetle (Dendroctonus ponderosae), have caused extensive timber losses in North America, with outbreaks in British Columbia alone projected to result in a cumulative GDP loss of $57.3 billion CAD and an annual GDP decrease exceeding 1.3% due to reduced wood supply and forest mortality.^[62] Similarly, weevils in the family Curculionidae, exemplified by the cotton boll weevil (Anthonomus grandis), have inflicted over $22 billion USD in yield losses and control costs on the U.S. cotton industry since their introduction in 1892, drastically reducing cotton acreage and necessitating ongoing eradication programs.^[63] On the beneficial side, certain Polyphaga species provide economic value in agriculture and beyond. Dung beetles (Scarabaeidae) enhance nutrient cycling in pastures by burying manure, which improves soil fertility, reduces parasite loads, and increases forage availability; in the UK, this service is estimated to save the cattle industry approximately £367 million annually through avoided losses in productivity and pest control.^[64] Ladybird beetles (Coccinellidae) serve as key biological control agents against aphids, with each individual in Chinese cotton fields providing at least 0.05 CNY (about 0.01 USD) in economic benefit by reducing pesticide needs; doubling their density can boost farmer incomes by roughly $100 USD per hectare per year.^[65] Additionally, carrion beetles (Silphidae) contribute to forensic entomology by aiding in post-mortem interval estimates during legal investigations, offering reliable evidence in criminal cases where traditional methods fail, thus supporting efficient judicial processes.^[66] Industrial applications of Polyphaga leverage their exoskeletons for chitin extraction, a biopolymer used in biomedical research, wound dressings, and sustainable materials. Chitin from beetle cuticles, such as those of Scarabaeidae species like Holotrichia spp., exhibits high purity and biocompatibility, enabling applications in drug delivery and tissue engineering with lower environmental impact than crustacean-derived sources.^[67]^[68]

Conservation status

Polyphaga, the largest suborder of beetles encompassing over 300,000 species, faces significant threats to its biodiversity primarily from habitat loss, climate change, and pesticide use. Habitat destruction through deforestation is a major driver, particularly in tropical regions where many Polyphagans are endemic and reliant on intact forest ecosystems; for instance, conversion of tropical dry forests to agriculture has led to substantial declines in coleopteran assemblages, including Polyphaga taxa like rove beetles (Staphylinidae).^[69]^[70] Climate change exacerbates these pressures by altering species distributions, with evidence of upward and poleward range shifts in European dung beetles (Scarabaeoidea), a key Polyphaga superfamily, as warmer temperatures enable colonization of higher elevations and northern latitudes while potentially contracting southern ranges.^[71] Additionally, pesticides pose risks to non-target Polyphagans, as broad-spectrum applications reduce populations of beneficial species like ground beetles (Carabidae) and leaf beetles (Chrysomelidae) through direct toxicity and sublethal effects on reproduction and foraging.^[72] Assessments on the IUCN Red List highlight the vulnerability of assessed Polyphaga species, with approximately 11-14% classified as threatened across subgroups such as saproxylic beetles and fireflies (Lampyridae); for example, in Europe, the proportion of threatened saproxylic Polyphagans ranges from 13.5% to 37.9% depending on data deficiencies (mid-point estimate 17.9% as of 2018).^[73]^[74]^[75] Specific cases underscore this, including endangered flightless stag beetles of the genus Apterocyclus on Kauai, Hawaii, which are critically imperiled due to habitat degradation and are among the few native Polyphagans in the archipelago.^[76] Polyphagans also serve as indicators of ecosystem health, with groups like dung beetles reflecting habitat quality through their sensitivity to soil degradation and fragmentation, aiding in broader biodiversity monitoring. As of 2025, IUCN updates indicate ongoing increases in insect threat levels due to habitat loss, though specific Polyphaga reassessments remain limited.^[77]^[78]^[79] Conservation efforts for Polyphaga focus on mitigating these threats through targeted strategies. Protected areas are crucial for safeguarding endemic hotspots, such as Mediterranean wetlands and Central Highland forests in Madagascar, where they preserve diverse aquatic and terrestrial Polyphagan communities, including water beetles (Hydrophiloidea).^[80]^[81] Integrated pest management (IPM) programs reduce reliance on broad-spectrum insecticides, promoting biological controls that spare non-target Polyphagans and maintain ecological balance in agricultural landscapes.^[82] Furthermore, citizen science initiatives, such as iNaturalist observations and dedicated apps like BOB (Behavioural Observations in Beetles), enable tracking of rare Polyphagans, including stag and longhorn beetles, to inform population monitoring and habitat restoration.^[83]^[84]

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The eggs of beetles are usually ovoid in shape with a plain surface, rarely exhibiting any marked sculpture, or diversity of form, such as we find in the ...<|control11|><|separator|>
[27]
[PDF] Rove Beetles of Florida, Staphylinidae (Insecta: Coleoptera
Eggs of Staphylinidae typically are white, spherical, spheroidal, or pyriform (pear-shaped). Almost all larvae are elongate and campodeiform, their antennae ...
[28]
Morphology of the abdominal segmental glands and spinning ...
The L3 larvae of both species have a campodeiform and elongated body shape (Fig. 1, Fig. 2A–B). Their heads are prognathous with six stemmata on each side and ...
[29]
Elateridae - an overview | ScienceDirect Topics
A horn-like terminal process known as urogomphi, surrounding a caudal notch, is present on the ninth abdominal segment. ... 5 Wireworms (Elateridae). The status ...
[30]
Click Beetles (Elaterids) - Missouri Department of Conservation
The larvae are thin, wormlike, shiny, hard-bodied, and segmented. Called wireworms, they have 6 tiny legs and are tan, whitish, or brownish. They look a bit ...
[31]
Holometabola: The Endopterygote Group - ENT 425
These insects have four developmental stages in the life cycle: egg, larva, pupa, and adult (imago). The larval stage is a period of active feeding and growth.
[32]
The stages of the beetle life cycle - Terminix
Mar 26, 2025 · Diapause can occur at each life stage of a beetle, helping ensure the species' survival. Pupae undergo diapause, pausing their transformation ...Missing: Polyphaga | Show results with:Polyphaga
[33]
Pachyschelus - Bugs With Mike
Reproductive habits are not specified; typical reproduction in beetles includes internal fertilization with oviposition of eggs. Predators. Information on ...
[34]
Effect of Soil Moisture on Plectris aliena (Coleoptera: Scarabaeidae ...
Sep 1, 2016 · This result is similar to observations in the current study, in which oviposition was greatest at the higher soil moisture levels (11% and 20%).
[35]
Effect of Soil Moisture and Soil Texture on Oviposition by Japanese ...
In nonchoice tests, Japanese beetles laid similar numbers of eggs into a wide range of soil textures, except fewer eggs were laid into pure sand. In choice ...
[36]
predation and the dispersal of larvae from the maternal burrow of ...
Females of the intertidal subsocial beetle, Bledius spectabilis Kraatz (Staphylinidae), look after their eggs and young larvae in burrows in saltmarsh mud.
[37]
(PDF) Staphylinidae and fungi - ResearchGate
Aug 22, 2015 · SETSUDA (1994) has reported subsocial parental care in the Oxyporinae. Adult females of O. japonicus Sharp have been observed to construct ...
[38]
[PDF] Asian Longhorned Beetle - Utah State University Extension
ALB's life cycle (Fig. 10) lasts from 1 to 3 years, and in northern China, a single generation takes 2 years (CABI, 2021).<|control11|><|separator|>
[39]
Coleoptera (Beetles) in Aquatic Ecosystems - ResearchGate
The suborder Polyphaga contains the majority of the beetle families. Polyphaga feeding habits range from predators (Hydrophilidae), to grazers on algae and ...
[40]
Diversity and distribution of polyphagan water beetles (Coleoptera ...
Feb 14, 2017 · Like the hydradephagans, polyphagans are also found in all types of aquatic habitats and although they do not spread into the open ocean, some ...
[41]
An Introduction to Ground Beetles: Beneficial Predators on Your Farm
Apr 8, 2020 · The group is distributed across North America and found in a wide range of habitats including grasslands, open meadows, forests, agricultural ...
[42]
Blister Beetles - UK Entomology - University of Kentucky
Blister beetles are members of a family of plant-feeding insects (Meloidae) that contain cantharidin, a toxic defensive chemical that protects them from ...
[43]
Wasp beetle | The Wildlife Trusts
A clever mimic, the wasp beetle is black-and-yellow and moves in a jerky, flight-like fashion - fooling predators into thinking it is actually a more harmful ...
[44]
Beetle-Fungus Interactions - Ambrosia Symbiosis
Symbiotic interactions between ambrosia beetles and ambrosia fungi are fascinating and there is a lot remaining to learn about them.
[45]
[PDF] Mycangia Define the Diverse Ambrosia Beetle– Fungus Symbioses
Mar 8, 2022 · This obligate mutualism (ambrosia symbiosis) allows both beetle and fungus to carve a unique niche in a nutrient- poor substrate (Hubbard 1897a, ...
[46]
Polyphaga - an overview | ScienceDirect Topics
The order Coleoptera is usually divided into four suborders ... All other beetles—150 families and 90% of the 340,000 species—are classified as Polyphaga.
[47]
New approaches narrow global species estimates for beetles ...
Jun 1, 2015 · Beetles account for roughly 25% (350,000–400,000 species) (1) of all described species (∼1.5 million species), making this the most species-rich ...Number Of Beetle Species In... · Results · Discussion
[48]
Transcriptomic data recover a new superfamily-level phylogeny of ...
The infraorder Cucujiformia, containing greater than 173,000 species in approximately 92 families worldwide, is the largest taxon in the suborder Polyphaga ...
[49]
Phylogeny and evolution of Staphyliniformia and Scarabaeiformia ...
Aug 6, 2014 · The beetle series Staphyliniformia exhibits extraordinary taxonomic, ecological and morphological diversity. To gain further insight into ...
[50]
Sampling and curation of rove beetles (Insecta, Coleoptera ...
Dec 19, 2022 · Introduction. With more than 64 000 described species, Staphylinidae (Insecta, Coleoptera), the rove beetles, is the second largest family ...
[51]
Chrysomelidae - an overview | ScienceDirect Topics
III.7. Chrysomelidae. The leaf beetles (35,000 species) are by far the largest group of beetles that primarily feed on the leaves of plants.
[52]
[PDF] Most diverse, most neglected: weevils (Coleoptera: Curculionoidea ...
May 25, 2023 · Weevils contain over 62,000 described species and a colossal amount of unknown diversity (Oberprieler et al., 2007). This beetle clade is almost ...
[53]
Staphylinidae - an overview | ScienceDirect Topics
The staphylinids are the least well known family of beetles, with perhaps only 10% of extant species described, and may eventually prove to be the largest ...
[54]
Phylogenomics of darkling beetles (Coleoptera: Tenebrionidae ...
Feb 23, 2023 · The complete mitochondrial genomes of four lagriine species (Coleoptera, Tenebrionidae) and phylogenetic relationships within Tenebrionidae.
[55]
Family BUPRESTIDAE Leach, 1815 - Australian Plant Census
The family Buprestidae is the eighth largest family of beetles in the world with approximately 14,700 species found in all biogeographical regions from the ...
[56]
The evolution and genomic basis of beetle diversity - PMC - NIH
Nov 18, 2019 · Beetle diversity thus appears to have resulted from multiple factors, including low extinction rates over a long evolutionary history ...
[57]
Origin and Diversification of Dung Beetles in Madagascar - PMC
Apr 20, 2011 · Dung beetles are a great example of Madagascar's biological uniqueness: of the almost 300 species described to date, 285 (96%) are endemic, with ...
[58]
A Jewel of the Pollinator World
May 19, 2025 · Jewel Beetles are in the Buprestidae family which is the eighth largest of the beetle families in the world. Nearly all species of this family ...Missing: buprestid | Show results with:buprestid
[59]
(PDF) Cretaceous West Gondwana vicariance shaped giant water ...
Aim We tested the hypothesis that ancient vicariance in giant water scavenger beetles shaped their current distribution. Location Worldwide except Antarctica.
[60]
Increasing global risk of khapra beetle invasion forecasted under ...
Jul 18, 2025 · Trogoderma granarium has been listed among the top 100 worst invasive species because of its damage, population growthand persistence once it ...
[61]
economic impact of the mountain pine beetle infestation in British ...
Nov 18, 2015 · The MPB infestation will cause a >1.3% annual GDP decrease, 1% income decline, and a $57.3 billion cumulative GDP loss, with $90 billion ...
[62]
Beating the $22 Billion Bug - USDA ARS
Feb 7, 2003 · The boll weevil wreaked havoc on the American cotton industry, with yield losses and control costs totaling more than $22 billion since its 1892 arrival in the ...
[63]
The application of an ecosystem services framework to estimate the ...
Aug 25, 2015 · We suggest that dung beetles may be currently saving the UK cattle industry c. £367 million each year: c. £354 million in conventional systems and c. £13 ...
[64]
Swapping pesticides for beetles could put money in farmers' pockets
Sep 11, 2018 · Cutting chemical pesticide use and relying on ladybird beetles to kill aphids can yield concrete benefits for Chinese cotton farmers.
[65]
Silphidae), a carrion beetle of forensic importance in the Palearctic ...
Jun 11, 2022 · Analysis of insects can provide evidence in death cases, for example, by answering the question about the time of death.
[66]
Extraction and Characterization of Chitin from the Beetle Holotrichia ...
The most common method for chitin extraction from insects involves two steps, an acidic step to remove catechols and a basic step to remove the cuticle proteins ...<|control11|><|separator|>
[67]
The Potential of Insects as Alternative Sources of Chitin - NIH
Jul 15, 2020 · The aim of this paper is to review the extraction method of chitin and chitosan for a comparison of the optimal demineralization and deproteinization processes.
[68]
Changes in Coleopteran assemblages over a successional ... - NIH
Jul 12, 2023 · In Latin America, the conversion of forest to agricultural, livestock fields, and urban centers has led to a loss of up to 25% of the original ...
[69]
Rove beetle species diversity and the patterns of interactions with ...
Oct 25, 2024 · Tropical rainforest deforestation and consequent habitat loss, usually distress plant and animal populations and the different types of biotic ...
[70]
Climate change and elevational range shifts: evidence from dung ...
Dec 20, 2013 · We investigate changes in the elevational distribution of dung beetle species (Coleoptera: Scarabaeoidea) in two separate mountain regions in Europe.Missing: scarab | Show results with:scarab
[71]
Pesticides have negative effects on non-target organisms - PubMed
Feb 13, 2025 · Pesticides affect a diverse range of non-target species and may be linked to global biodiversity loss. The magnitude of this hazard remains only partially ...
[72]
Habitat loss blamed for more species decline | IUCN
Mar 12, 2010 · Nine percent of butterflies, 11 percent of saproxylic beetles (beetles that depend on decaying wood) and 14 percent of dragonflies are threatened with ...
[73]
[PDF] EuropEan rEd List of saproxyLic BEEtLEs - IUCN Portals
Thus, the proportion of threatened species could lie between 13.5% (if all DD species are not threatened) and 37.9% (if all. DD species are threatened) for ...
[74]
Evaluating firefly extinction risk: Initial red list assessments for North ...
Nov 17, 2021 · We found at least 18 species (14%) are threatened with extinction (eg categorized as Critically Endangered, Endangered, or Vulnerable) due to various pressures.
[75]
Native Scarab Beetles in Hawaiʻi - CRB Hawaii
Oct 11, 2021 · These flightless, endemic beetles are known from Kauaʻi and are extremely rare. Native Hawaiian stag beetles, like many endemic species ...
[76]
(PDF) The IUCN Red List of Threatened Species - ResearchGate
Dec 24, 2020 · Of the 644 species of dung beetles inhabiting the Mediterranean region, 200 (32%) have at least 75% of their distribution range within the ...
[77]
How does landscape composition and configuration affect dung ...
Oct 3, 2025 · Because they are sensitive to environmental changes, dung beetles are often used as bioindicators of landscape health (Nichols et al. 2007 ...
[78]
A case study using species distribution models for water beetles in ...
Results showed that the highest potential species richness of water beetles in Morocco occurs in the mountain areas of the Rif and Prerif, Middle Atlas and ...
[79]
Remaining forests on the Central Highlands of Madagascar ...
Dec 12, 2022 · Here, we analyze the aquatic Adephaga beetle fauna of three remaining protected forests of the Central Highlands. Diversity, abundance, and ...
[80]
Integrated Pest Management (IPM) Principles | US EPA
Sep 2, 2025 · IPM is an effective and environmentally sensitive approach to pest management that relies on a combination of common-sense practices.Missing: Polyphaga | Show results with:Polyphaga
[81]
From Phenology and Habitat Preferences to Climate Change
Apr 20, 2021 · The aim of this paper is to evaluate the role of citizen science, particularly observations done using the iNaturalist website (iNaturalist.org) ...
[82]
Final Release of the BOB Citizen Science App (Behavioural ...
May 6, 2025 · The BOB App not only encourages citizen scientists to celebrate finding a beetle but also to sit down, take their time, and carefully observe ...