Hubbry Logo
ScarabaeidaeScarabaeidaeMain
Open search
Scarabaeidae
Community hub
Scarabaeidae
logo
8 pages, 0 posts
0 subscribers
Be the first to start a discussion here.
Be the first to start a discussion here.
Scarabaeidae
Scarabaeidae
Scarabaeidae
View on Wikipedia
from Wikipedia

Scarab beetle
Central European scarab beetles
with some anatomical details. Edmund Reitter's Fauna Germanica, 1908
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Polyphaga
Infraorder: Scarabaeiformia
Superfamily: Scarabaeoidea
Family: Scarabaeidae
Latreille, 1802
Subfamilies[1][2][3]
On this high quality closeup, head anatomic details are well visible.

The family Scarabaeidae, as currently defined, consists of over 35,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change. Several groups formerly treated as subfamilies have been elevated to family rank (e.g., Bolboceratidae, Geotrupidae, Glaresidae, Glaphyridae, Hybosoridae, Ochodaeidae, and Pleocomidae), and some reduced to lower ranks. The subfamilies listed in this article are in accordance with those in Catalog of Life (2023).[3]

Description

[edit]
Sacred scarab in a cartouche of Thutmosis III from Karnak temple of Amun-Ra, Egypt

Scarabs are stout-bodied beetles; most are brown or black in colour, but many, generally species that are diurnally active, have bright metallic colours, measuring between 1.5 and 160 millimetres (0.059 and 6.3 in). The antennae of most species superficially seem to be knobbed (capitate), but the several segments comprising the head of the antenna are, as a rule, lamellate: they extend laterally into plates called lamellae that they usually keep compressed into a ball. Then, when following a scent, such a beetle fans the lamellae out like leaves to sense odours.

Many species are fossorial, with legs adapted for digging. In some groups males (and sometimes females) have prominent horns on the head and/or pronotum to fight over mates or resources.[4] The largest fossil scarabaeid was Oryctoantiquus borealis with a length of 50 millimetres (2.0 in).[5]

A scarab beetle grub from Australia.

The C-shaped larvae, called grubs, are pale yellow or white. Most adult beetles are nocturnal, although the flower chafers (Cetoniinae) and many leaf chafers (Rutelinae) are active during the day. The grubs mostly live underground or under debris, so are not exposed to sunlight. Many scarabs are scavengers that recycle dung, carrion, or decaying plant material.[6] Others, such as the Japanese beetle, are plant-eaters, wreaking havoc on various crops and vegetation.

Some of the well-known beetles from the Scarabaeidae are Japanese beetles, dung beetles, June beetles, rose chafers (Australian, European, and North American), rhinoceros beetles, Hercules beetles and Goliath beetles.

Several members of this family have structurally coloured shells which act as left-handed circular polarisers; this was the first-discovered example of circular polarization in nature.[7]

Ancient Egypt

[edit]

In Ancient Egypt, the dung beetle now known as Scarabaeus sacer (formerly Ateuchus sacer) was revered as sacred.[8] Egyptian amulets representing the sacred scarab beetles were traded throughout the Mediterranean world.[4]

See also

[edit]

References

[edit]

Further reading

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Scarabaeidae

View on Grokipedia
from Grokipedia
Scarabaeidae is a large and diverse family of beetles within the order Coleoptera, commonly known as scarab beetles, encompassing over 30,000 described species distributed worldwide. These insects are characterized by their robust, often oval-shaped bodies, lamellate (club-like) antennae that can be folded into a cup, and stout legs adapted for various lifestyles, including digging and climbing. Adults typically range in size from a few millimeters to over 10 centimeters, with the larval stages of some species, like the Goliath beetle (Goliathus goliathus), reaching up to 100 grams, ranking among the heaviest insect larvae. The family Scarabaeidae, established taxonomically by in 1802, belongs to the superfamily and includes numerous subfamilies such as Scarabaeinae (dung beetles), (chafers and June beetles), (rhinoceros and Hercules beetles), and Cetoniinae (flower chafers). This classification reflects their morphological and ecological diversity, with larvae often C-shaped, white grubs that inhabit soil, dung, or decaying wood, feeding on roots, , or . Scarabaeids exhibit complete , with life cycles varying from one to several years depending on the species and environment. Ecologically, Scarabaeidae play crucial roles in ecosystems, particularly through nutrient recycling, soil aeration, and secondary , especially in the coprophagous subfamilies that process animal dung. Many species are phytophagous, with adults consuming foliage, flowers, or sap, while some act as pollinators or pests in ; for instance, white grubs of the Phyllophaga damage turf and crops by feeding on roots. Their cosmopolitan distribution spans forests, grasslands, deserts, and agricultural lands, underscoring their adaptability and importance in maintaining and .

Taxonomy and Phylogeny

Classification

Scarabaeidae belongs to the order Coleoptera and the superfamily , comprising one of the most species-rich families within the beetles, with over 35,000 described species distributed across approximately 1,600 genera worldwide. This superfamily represents a major lineage in the phylogeny, encompassing diverse forms adapted to various ecological niches. Phylogenetically, Scarabaeidae is included in the series Scarabaeiformia, as defined by Crowson in , which groups it with other scarabaeoid lineages. Recent molecular studies, including transcriptome-based analyses, have confirmed close evolutionary relationships between Scarabaeidae and families such as Lucanidae (stag beetles) and (bess beetles), often placing them in a basal within alongside Geotrupidae. These findings support a revised understanding of scarabaeoid interfamily connections, highlighting shared ancestral traits like specialized antennal structures. The family was originally established by in 1802 as a broad assemblage of scarab-like beetles. Over time, taxonomic revisions based on morphological and molecular evidence have led to the elevation of several subfamilies to independent family status in contemporary classifications, including Geotrupidae (earth-boring dung beetles) and Hybosoridae (scavenger scarab beetles). At the family level, Scarabaeidae is distinguished by key diagnostic features in adults, notably the lamellate antennae—composed of plate-like segments that can fold into a club—and a transverse pygidial plate on the terminal abdominal segment. These traits aid in differentiating Scarabaeidae from closely related scarabaeoid families and underscore its monophyletic status in modern systematics.

Subfamilies and Diversity

The family Scarabaeidae encompasses approximately 28 subfamilies, representing one of the most taxonomically diverse groups within the Coleoptera order. This subdivision highlights the family's , with subfamilies exhibiting distinct morphological adaptations suited to varied ecological niches, from subterranean to arboreal lifestyles. The total exceeds 35,000, with approximately 36,000 described species as of 2025 and ongoing discoveries suggesting even higher actual diversity, particularly in understudied tropical regions where the majority of species occur. Among the major subfamilies, Scarabaeinae, commonly known as dung beetles, includes over 5,000 described species across more than 200 genera, with tribes such as Coprini featuring notable genera like , renowned for their specialized forms. , encompassing chafers and June beetles, stands as one of the largest subfamilies with approximately 11,000 species in over 750 genera, exemplifying the family's soil-dwelling adaptations. , or rhinoceros beetles, comprises robust species known for their impressive size and horn-like structures, contributing to the subfamily's estimated several thousand species. Cetoniinae, the flower chafers, features vibrant, metallic species adapted to floral resources, with genera such as overlapping in size extremes within the family. Rutelinae, including shining leaf chafers, harbors pests like Popillia japonica () and accounts for around 5,000 species, underscoring the subfamily's phytophagous tendencies. Aphodiinae, a group of smaller dung beetles, adds further diversity with species that often inhabit temperate and arid zones, complementing the more tropical dominance seen in other subfamilies. This subfamily structure illustrates the Scarabaeidae's global variability, with tropical hotspots driving much of the evolutionary innovation in form and habitat occupation.

Morphology and Physical Characteristics

Adult Features

Adult Scarabaeidae beetles exhibit a stout, compact body build, typically rounded or in shape, which contributes to their robust appearance. These beetles vary widely in size, ranging from approximately 2 mm in small species such as certain aphodiines (e.g., Aphodius spp.) to over 100 mm in large forms like the Goliath beetle (Goliathus goliathus). Characteristic features include lamellate antennae composed of 3 to 7 flattened segments forming a club used for sensory detection, which can be folded or fanned out. The legs are often clubbed or equipped with teeth, particularly on the protibiae, adaptations that facilitate digging, burrowing, or climbing in various habitats. Many species display metallic or iridescent coloration, such as the vibrant green hues of the June beetle (), resulting from structural properties in the . The head is typically deflexed, with forward-projecting mandibles that are often lamelliform and adapted for feeding on diverse substrates. The pygidium, the terminal abdominal tergite, is frequently exposed beyond the elytra and transverse in orientation, a diagnostic trait visible in many taxa. Sexual dimorphism is prominent in several subfamilies, notably , where males develop exaggerated horns on the head or pronotum, while females lack such structures; for instance, in Dynastes hercules, males possess prominent cephalic and thoracic horns.

Larval Characteristics

Larvae of the Scarabaeidae, commonly referred to as white grubs, display a characteristic C-shaped body when at rest, with a creamy-white and a hardened, brown head capsule. These larvae possess three pairs of prominent thoracic legs adapted for movement within environments. The body surface is often adorned with fine setae or fringes that assist in burrowing through substrate. Notably, Scarabaeidae larvae lack urogomphi, the caudal spines present in larvae of some other scarabaeoid families, which further distinguishes their morphology. The head capsule houses well-developed, sclerotized mandibles suited for chewing such as roots or . Larvae generally progress through three s, with sizes increasing markedly from about 5 in the first instar to over 50 in the mature third instar, varying by and environmental conditions. A prominent diagnostic feature is the raster, the patterned arrangement of spines and setae on the ventral anal plate of the terminal abdominal segment, which varies across subfamilies and enables taxonomic identification. For instance, larvae typically exhibit a Y-shaped anal slit within the raster, often accompanied by distinctive rows of short spines. In contrast, Scarabaeinae grubs feature transverse anal slits, with the raster showing parallel or clustered setae patterns adapted to their detritivorous habits.

Life Cycle and Biology

Reproduction and Development

Reproduction in Scarabaeidae begins with adult emergence, which occurs either diurnally or nocturnally depending on the . Males typically attract females through the release of sex pheromones or via physical displays, including combative horn fights observed in the subfamily to secure rights. Following copulation, females oviposit 20 to 100 eggs, often in batches, directly into soil, dung pats, or decaying plant matter to provide suitable conditions for early development. Scarabaeidae eggs are generally pearl-like, white or translucent, and measure about 1-2 in diameter; they are laid either singly or in small clusters within prepared burrows or chambers. The typically ranges from 1 to 2 weeks, during which embryos develop under favorable temperatures around 20-30°C. Upon hatching, larvae progress through three distinct s, a characteristic feature of the family Scarabaeidae. The larval stage is the longest in the life cycle, enduring 1 to 4 years overall—for instance, about 3 years in June beetles of the genus —during which the grubs grow substantially and often enter to overwinter. Pupation follows in the final , where the mature constructs an earthen cell in the soil for , a process lasting 2 to 6 weeks depending on temperature. The total life cycle from to emergent spans 1 to 4 years, influenced markedly by species-specific traits and climatic variables. Environmental factors such as and play critical roles, with optimal ranges (e.g., 15-25°C and moderate ) promoting continuous development, while extremes trigger in larvae to enhance survival through adverse seasons.

Feeding and Behavior

Adult scarab beetles exhibit diverse feeding strategies depending on the subfamily. Members of the Scarabaeinae primarily consume dung from mammalian herbivores, using specialized mouthparts to manipulate this soft substrate. In contrast, adults of the Cetoniinae feed on and from flowers, often acting as pollinators during diurnal foraging. Melolonthinae adults typically chew foliage from trees and shrubs, though some species also consume flowers or . Certain scarab adults, such as those in the Pleocominae, do not feed at all, relying on stored energy reserves from the larval stage. Larvae of Scarabaeidae, commonly known as white grubs in many species, primarily feed on plant roots, , or dung in the . Early instars often consume and decaying material, transitioning to root-feeding in later stages, which can damage turf and crops. For instance, white grubs of species burrow and feed on grass roots near the surface. Behavioral patterns in Scarabaeidae are adapted to their feeding ecologies. Dung-rolling species, such as those in the genus within Scarabaeinae, form balls of dung and roll them away from the source using straight paths guided by celestial cues like the sun, , and . Other dung beetles employ tunneling strategies, burying dung underground, or dwelling within it, to secure food resources. is observed in some species, where adults bury dung provisions for , with biparental in nest maintenance and guarding. Sensory adaptations facilitate these behaviors, particularly through the antennae, which bear lamellae equipped with sensilla for detecting volatile organic compounds like odors from dung or nectar. Pore plates on the antennal lamellae house numerous olfactory neurons, enabling precise localization of food sources. Many scarabs, including June beetles (Phyllophaga spp.) of the Melolonthinae, exhibit nocturnal activity, emerging at dusk to feed and mate while avoiding diurnal predators.

Distribution and Ecology

Global Range

The family Scarabaeidae exhibits a , occurring on all continents except and being absent from oceanic environments. With over 30,000 worldwide, the family demonstrates highest in tropical regions, where environmental conditions support a wide array of subfamilies and genera. Regionally, stands out as a hotspot for dung beetles (subfamily ), hosting the greatest global diversity of these due to abundant mammalian herbivores providing dung resources. In , rhinoceros beetles (subfamily ) are particularly dominant, with many adapted to tropical and temperate forests across and . has seen significant introductions of Scarabaeidae, exemplified by the (Popillia japonica), which was accidentally brought from to the in 1916 via nursery stock and has since become a widespread invasive pest east of the . Dispersal of Scarabaeidae occurs naturally through flight, wind currents, and attachment to animals, enabling of new areas over evolutionary timescales. Human-mediated dispersal has accelerated range expansions, including intentional releases for biological control and unintentional transport via trade, as seen with Digitonthophagus gazella introductions to the and ongoing spread of P. japonica. Endemism is pronounced in isolated regions like , where nearly all species (Scarabaeinae) are endemic, reflecting long-term evolutionary isolation. However, habitat loss from and land-use changes poses a major threat, contracting ranges and reducing population viability for many endemic and tropical species.

Habitats and Ecological Roles

Scarabaeidae species inhabit diverse environments worldwide, primarily associated with in grasslands, , deserts, and agricultural fields, where many dung-feeding members thrive on feces. Some subfamilies, such as Cetoniinae (flower chafers), are arboreal and frequent canopies or flowering , while others occupy riparian zones along rivers and streams bordered by native vegetation. These beetles play critical roles in ecosystem processes, particularly through dung decomposition by Scarabaeinae species, which bury substantial amounts of manure—up to approximately 2 metric tons per hectare per year in some pastoral systems—accelerating breakdown and reducing parasite transmission to livestock and wildlife. By tunneling into soil, they enhance aeration, improving water infiltration and root penetration, while facilitating nutrient cycling through the release of nitrogen and phosphorus from buried dung, thereby boosting soil fertility and plant growth. Additionally, dung beetles contribute to secondary seed dispersal by transporting and burying seeds within fecal pats, promoting germination away from parent plants and reducing predation risk. Flower chafers aid in secondary pollination by feeding on nectar and pollen in flowers, transferring pollen between plants during diurnal activity. Ecological interactions include serving as prey for birds, mammals, and other arthropods, which helps regulate populations and supports higher trophic levels. They also compete with other decomposers like for dung resources, influencing microbial communities and breakdown rates. Scarabaeidae assemblages are sensitive indicators of , with their diversity and abundance reflecting habitat quality and metrics in monitoring programs. Conservation concerns arise from population declines driven by pesticide use, which directly kills non-target individuals, and habitat fragmentation, which disrupts dung availability and connectivity, thereby impairing their roles in decomposition and nutrient cycling.

Interactions with Humans

Economic Importance

Scarabaeidae exhibit a dual economic role in agriculture, serving as significant pests that inflict substantial damage to crops and turf while also providing valuable ecosystem services through species like dung beetles that enhance soil fertility and reduce pest pressures in livestock systems. Larval stages, known as white grubs, feed on plant roots, causing severe damage to turfgrass, lawns, and field crops such as corn and soybeans, with affected areas often wilting and dying due to disrupted water and nutrient uptake. For instance, the Japanese beetle (Popillia japonica), a notorious invasive scarab, has larvae that alone contribute to annual economic losses exceeding $450 million in the United States through damage to turf and ornamental plants, compounded by control costs that push the total impact over $460 million yearly (estimated in the early 2000s). As of 2025, the Japanese beetle's spread to new areas, such as increased populations in Washington state, continues to amplify economic concerns. Adult scarabs exacerbate these losses by defoliating foliage, flowers, and fruits; June beetles (Phyllophaga spp.), for example, can strip leaves from orchard trees like apples and pecans at night, leading to reduced photosynthesis and yield declines that threaten fruit production, though precise national figures are integrated into broader scarab pest valuations. Other examples include rhinoceros beetles (Oryctes rhinoceros), which bore into the crowns of oil palm and coconut trees, causing up to 25% annual crop losses in palm oil plantations and 40-92% yield reductions in young replants, severely impacting global tropical agriculture. Similarly, chafer beetles such as the rose chafer (Cetonia aurata) damage vineyards by skeletonizing grape leaves and flowers, with economic thresholds as low as two adults per vine triggering interventions to prevent blossom loss and reduced berry set in wine production. In contrast, many Scarabaeidae, particularly dung beetles in the subfamilies Scarabaeinae and Aphodiinae, deliver substantial economic benefits by recycling manure, which improves pasture productivity and mitigates in systems. These beetles bury and decompose up to 80-90% of dung pats, incorporating nutrients like back into the and enhancing growth by 20-30% in treated pastures, while also suppressing populations of pest flies and parasites that affect health. In the United States, the ecosystem services from s— including nutrient cycling, reduced fly breeding, and decreased parasite loads—are valued at approximately $530 million as of 2023 for operations, averting higher costs in feed, veterinary care, and land maintenance. Introduced dung beetle species in , such as Onthophagus and Euoniticellus taxa, have similarly bolstered biological control since the , burying dung to cut bush fly numbers by up to 70%, recycle nutrients for better , and generate ongoing economic gains estimated at over $1 billion annually through improved pasture sustainability and reduced chemical inputs. Management of pestiferous Scarabaeidae relies on (IPM) strategies that balance chemical, cultural, and biological controls to minimize economic losses while preserving beneficial . Biological options include applying entomopathogenic nematodes (Heterorhabditis bacteriophora) to target soil-dwelling grubs, which parasitize and kill larvae with efficacy rates of 50-90% under optimal moist conditions, and milky spore disease (Paenibacillus popilliae), a bacterium that infects grubs specifically and can provide suppression for up to 10-20 years after initial application, though effectiveness varies according to recent studies. Additionally, essential oils such as clove, cinnamon, thyme, and basil have demonstrated toxic effects on Scarabaeidae beetle larvae, including growth inhibition and mortality exceeding 90% in some cases, and are employed in controlling pest species like the coconut rhinoceros beetle (Oryctes rhinoceros). Cultural practices, such as and endophyte-enhanced grasses, further disrupt grub habitats, while economic assessments highlight the value of conserving dung beetles, whose services offset pest management costs in mixed agroecosystems.

Cultural and Symbolic Significance

In , scarab beetles, particularly , held profound symbolic importance as embodiments of rebirth, creation, and the sun's daily cycle, closely associated with the god , who was depicted with a beetle head and represented the rising sun pushing the solar disk across the sky. This symbolism stemmed from observations of the beetle rolling dung balls, interpreted as mimicking the sun's movement and the emergence of new life from the buried brood, thus linking the insect to renewal and immortality. Sacred scarabs were extensively used in artifacts, including amulets for protection and good fortune, jewelry such as rings and pendants, and administrative seals bearing royal names or hieroglyphs. Heart scarabs, placed over the deceased's heart during mummification, were inscribed with Spell 30B from the Book of the Dead to prevent the heart from testifying against the owner in the afterlife judgment before Osiris, ensuring a favorable outcome for eternal life. Thousands of such scarab artifacts, including over 10,000 amulets and seals, have been discovered in tombs and settlements, underscoring their ubiquity across social classes from the Middle Kingdom onward. Beyond , scarab beetles played minor roles in other cultures, often as symbols of transformation. Some Native American legends feature beetles as symbols of transformation and , though not specifically scarabs, and less prominently than in Egyptian lore; roles in broader Asian or sub-Saharan African traditions remain rare and undocumented beyond superficial motifs. In contemporary contexts, scarabs retain symbolic value through modern jewelry and tattoos, where designs evoke Egyptian motifs of and renewal, popular among those seeking talismans for personal growth. Media depictions, notably in films like The Mummy (1999), portray scarabs as voracious, flesh-burrowing horrors, contrasting their historical reverence and amplifying public fascination with ancient Egyptian heritage. This cultural legacy aids conservation efforts, as artifacts and symbolism raise awareness for threatened scarab species facing loss, integrating entomological with Egyptological studies. Post-Egyptian reverence waned after the Greco-Roman adoption of scarabs as mere luck charms, yet surviving artifacts continue to shape modern interpretations of ancient beliefs.

References

  1. https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/scarabaeidae
  2. https://genent.cals.ncsu.edu/insect-identification/order-coleoptera/family-scarabaeidae/
  3. https://www.britannica.com/animal/African-goliath-beetle
  4. https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=926326
  5. https://edis.ifas.ufl.edu/publication/IN202
  6. https://faculty.ucr.edu/~legneref/identify/scarabae.htm
  7. https://pmc.ncbi.nlm.nih.gov/articles/PMC3583983/
  8. https://soil.evs.buffalo.edu/index.php/Scarabaeidae
  9. https://pmc.ncbi.nlm.nih.gov/articles/PMC11153540/
  10. https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1120&context=entomologypapers
  11. https://resjournals.onlinelibrary.wiley.com/doi/10.1111/syen.12602
  12. https://www.gbif.org/species/112781435
  13. https://unsm-ento.unl.edu/Guide/Scarabaeoidea/Hybosoridae/Hybosoridae-Overview/HybosoridaeO.html
  14. https://unsm-ento.unl.edu/Guide/Scarabaeoidea/Scarabaeoidea-pages/Scarabaeoidea-Overview/ScarabaeoideaO.html
  15. https://pmc.ncbi.nlm.nih.gov/articles/PMC4829968/
  16. https://zookeys.pensoft.net/article/4001/
  17. https://www.mapress.com/mt/article/view/megataxa.12.1.1/53589
  18. https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2023.1168754/full
  19. https://unsm-ento.unl.edu/Guide/Scarabaeoidea/Scarabaeidae/Scarabaeinae/Scarabaeinae-Overview/ScarabaeinaeO.html
  20. https://www.entomoljournal.com/archives/2016/vol4issue5/PartN/4-5-103-628.pdf
  21. https://unsm-ento.unl.edu/Guide/Scarabaeoidea/Scarabaeidae/Scarabaeidae-pages/Scarabaeidae-Overview/ScarabaeidaeO.html
  22. https://www.sciencedirect.com/science/article/abs/pii/S1226861518302164
  23. https://bugguide.net/node/view/187
  24. https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2022.1008477/full
  25. https://www.popillia.eu/about-the-japanese-beetle-popillia-japonica/the-ipm-popillia-field-guide-to-the-most-common-european-scarab-beetles
  26. https://bugswithmike.com/factsheet/scarabaeidae
  27. https://ipm.ucanr.edu/PMG/GARDEN/FRUIT/PESTS/grfruitbeetle.html
  28. https://content.ces.ncsu.edu/rainbow-scarab-1
  29. https://www.ndsu.edu/faculty/rider/Schafer_Post/PDFs/007a.pdf
  30. https://edis.ifas.ufl.edu/publication/IN1142
  31. https://zookeys.pensoft.net/article/29236/
  32. https://www.ideals.illinois.edu/items/26809/bitstreams/91552/data.pdf
  33. https://www.sciencedirect.com/science/article/abs/pii/S0044523120300899
  34. https://www.scholarsresearchlibrary.com/articles/diversity-and-characterisation-of-phytophagous-scarabaeids.pdf
  35. https://www.sciencedirect.com/science/article/pii/S0960982223017578
  36. https://stri-apps.si.edu/docs/publications/pdfs/1987_JKES_Use_of_horns_in_flights_by_the_dimorphic_males_of_Ageopsis_nigricollis.pdf
  37. https://portal.ct.gov/caes/fact-sheets/entomology/the-japanese-beetle-popillia-japonica-newman-scarabaeidae-coleoptera
  38. https://preprints.arphahub.com/article/154400/download/pdf/
  39. https://www.cambridge.org/core/journals/bulletin-of-entomological-research/article/egg-development-and-viability-in-three-species-of-cyclocephala-coleoptera-scarabaeidae-dynastinae/B9AECC6B2BD2FEBE052F10AA3D26E52F
  40. https://pubmed.ncbi.nlm.nih.gov/23448133/
  41. https://www.researchgate.net/publication/234149637_Biology_of_Scarabaeidae
  42. https://edis.ifas.ufl.edu/publication/IN630
  43. https://www.tandfonline.com/doi/full/10.1080/03014223.2018.1545679
  44. https://resjournals.onlinelibrary.wiley.com/doi/10.1111/j.1365-3032.2008.00640.x
  45. https://webdoc.agsci.colostate.edu/bspm/JapaneseBeetle/PotterHeld2002.pdf
  46. https://resjournals.onlinelibrary.wiley.com/doi/10.1046/j.1365-2311.2002.00399.x
  47. https://www.researchgate.net/publication/262694310_A_review_of_the_natural_history_of_adult_Cetoniinae_Coleoptera_Scarabaeidae_from_Argentina_and_adjacent_countries
  48. https://ir.library.oregonstate.edu/downloads/t148fm82v
  49. https://pmc.ncbi.nlm.nih.gov/articles/PMC4802167/
  50. https://extension.okstate.edu/programs/turfgrass-science/educational-materials/white-grubs.html
  51. https://pmc.ncbi.nlm.nih.gov/articles/PMC3886324/
  52. https://www.sciencedirect.com/science/article/pii/S0960982212015072
  53. https://www.researchgate.net/publication/229800307_The_evolution_of_parental_behaviour_in_Scarabaeinae_Coleoptera_Scarabaeidae_an_experimental_approach
  54. https://www.sciencedirect.com/science/article/pii/S0003347202930368
  55. https://www.frontiersin.org/journals/ecology-and-evolution/articles/10.3389/fevo.2021.759778/full
  56. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0041357
  57. https://mdc.mo.gov/discover-nature/field-guide/may-beetles-june-bugs
  58. https://www.plu.edu/biology/wp-content/uploads/sites/7/2017/09/final_scarabaeidae_20170914.pdf
  59. https://www.britannica.com/animal/scarab-beetle
  60. https://www.mdpi.com/1424-2818/15/10/1066
  61. https://www.jstage.jst.go.jp/article/ggs/91/4/91_16-00003/_html/-char/en
  62. https://www.invasivespeciesinfo.gov/terrestrial/invertebrates/japanese-beetle
  63. https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1655&context=entomologyfacpub
  64. https://link.springer.com/article/10.1007/s10340-023-01653-1
  65. https://pmc.ncbi.nlm.nih.gov/articles/PMC4553453/
  66. https://besjournals.onlinelibrary.wiley.com/doi/10.1002/2688-8319.12297
  67. https://uwm.edu/field-station/bug-of-the-week/dung-beetle/
  68. https://www.researchgate.net/publication/371060713_Spatio-temporal_variation_of_dung_beetle_Coleoptera_Scarabaeidae_assemblages_in_a_community_ecological_reserve_of_southeastern_Mexico
  69. https://www.eolss.net/sample-chapters/C20/E6-142-TPE-02.pdf
  70. https://www.sciencedirect.com/science/article/pii/S0308521X21002390
  71. https://www.sciencedirect.com/science/article/abs/pii/S1146609X25000736
  72. https://www.researchgate.net/publication/222553331_Ecological_functions_and_ecosystem_services_provided_by_Scarabaeinae_dung_beetles
  73. https://zookeys.pensoft.net/article/60036/
  74. https://www.researchgate.net/publication/379989298_Declining_Dung_Beetle_Coleoptera_Scarabaeidae_Abundance_and_Diversity_in_the_Neotropics_Causes_and_Conservation_Strategies
  75. https://pmc.ncbi.nlm.nih.gov/articles/PMC10451379/
  76. https://extension.psu.edu/japanese-beetle-control-on-ornamental-plants/
  77. https://washingtonstatestandard.com/2025/10/02/more-money-needed-for-fight-against-two-invasive-bugs-wa-agriculture-department-says/
  78. https://citybugs.tamu.edu/2015/05/14/may-beetles-on-pecans/
  79. https://storymaps.arcgis.com/stories/32c7caf33e664eb09622290c2fcd3d69
  80. https://tuengr.com/V12/12A9S.pdf
  81. https://fruit.wisc.edu/2021/06/21/managing-japanese-beetle-and-rose-chafer/
  82. https://academic.oup.com/bioscience/article/56/4/311/229003
  83. https://www.townandcountrytoday.com/athabasca-news/dung-beetles-provide-millions-of-dollars-in-economic-benefits-6809181
  84. https://www.abc.net.au/news/rural/2018-09-13/dung-beetles-produce-economic-turnover-for-agriculture-industry/10231166
  85. https://www.sciencedirect.com/science/article/abs/pii/S1439179117300336
  86. https://extension.umd.edu/resource/white-grub-management-lawns
  87. https://www.fairfaxgardening.org/wp-content/webdocs/pdf/MilkySpore.pdf
  88. https://hgic.clemson.edu/factsheet/white-grub-management-in-turfgrass/
  89. https://gms.ctahr.hawaii.edu/gs/handler/getmedia.ashx?moid=72372&dt=3&g=12
  90. https://pubmed.ncbi.nlm.nih.gov/23448028/
  91. https://www.researchgate.net/publication/283101845
  92. https://isac.uchicago.edu/content/khepri-my-scarab
  93. https://unsm-ento.unl.edu/Egyptian_Sacred_Scarab/egs-text.htm
  94. https://clas.wayne.edu/biosci/news/how-the-scarab-joined-the-club-61959
  95. https://mcclungmuseum.utk.edu/1996/01/01/sacred-scarab/
  96. https://hoodmuseum.dartmouth.edu/objects/13.157.4112
  97. https://www.wheaton.edu/about-wheaton/museums-library-and-collections/archaeology-museum/thutmoses-iii-scarab/
  98. https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1094&context=nebanthro
  99. http://www.native-languages.org/legends-beetle.htm
  100. https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1093&context=entomologypapers
  101. https://a-z-animals.com/animals/beetle/beetle-facts/flesh-eating-beetles/
  102. https://www.nationalgeographic.com/animals/invertebrates/facts/scarabs
  103. https://www.researchgate.net/publication/228384734_Scarab_Beetles_in_Human_Culture
Add your contribution
Related Hubs
User Avatar
No comments yet.