
- •Contents
- •Foreword
- •How to use this book
- •Advisory boards
- •Contributing writers
- •Contributing illustrators
- •What is an insect?
- •Evolution and systematics
- •Structure and function
- •Life history and reproduction
- •Ecology
- •Distribution and biogeography
- •Behavior
- •Social insects
- •Insects and humans
- •Conservation
- •Protura
- •Species accounts
- •Collembola
- •Species accounts
- •Diplura
- •Species accounts
- •Microcoryphia
- •Species accounts
- •Thysanura
- •Species accounts
- •Ephemeroptera
- •Species accounts
- •Odonata
- •Species accounts
- •Plecoptera
- •Species accounts
- •Blattodea
- •Species accounts
- •Isoptera
- •Species accounts
- •Mantodea
- •Species accounts
- •Grylloblattodea
- •Species accounts
- •Dermaptera
- •Species accounts
- •Orthoptera
- •Species accounts
- •Mantophasmatodea
- •Phasmida
- •Species accounts
- •Embioptera
- •Species accounts
- •Zoraptera
- •Species accounts
- •Psocoptera
- •Species accounts
- •Phthiraptera
- •Species accounts
- •Hemiptera
- •Species accounts
- •Thysanoptera
- •Species accounts
- •Megaloptera
- •Species accounts
- •Raphidioptera
- •Species accounts
- •Neuroptera
- •Species accounts
- •Coleoptera
- •Species accounts
- •Strepsiptera
- •Species accounts
- •Mecoptera
- •Species accounts
- •Siphonaptera
- •Species accounts
- •Diptera
- •Species accounts
- •Trichoptera
- •Species accounts
- •Lepidoptera
- •Species accounts
- •Hymenoptera
- •Species accounts
- •For further reading
- •Organizations
- •Contributors to the first edition
- •Glossary
- •Insects family list
- •A brief geologic history of animal life
- •Index

Vol. 3: Insects |
Order: Coleoptera |
Species accounts
Giraffe-necked weevil
Trachelophorus giraffa
FAMILY
Attelabidae
TAXONOMY
Aploderus (Trachelophorus) giraffa Jekel, 1860, “Madagascar.”
OTHER COMMON NAMES
English: Red-and-black giraffe beetle; French: Scarabée girafe; Dutch: Giraf nek kever; German: Giraffenrüssler.
PHYSICAL CHARACTERISTICS
Males up to 0.98 in (2.5 cm). Black with red elytra; only males have long “neck.”
DISTRIBUTION
Madagascar.
HABITAT
Forests.
SIGNIFICANCE TO HUMANS
None known.
Giant metallic ceiba borer
Euchroma gigantea
FAMILY
Buprestidae
TAXONOMY
Buprestis gigantea Linnaeus, 1758, “America.”
OTHER COMMON NAMES
Spanish: Eucroma, catzo; Portuguese: Mae do sol, ôlho do sol.
PHYSICAL CHARACTERISTICS
Grows to 2–2.8 in (5–7 cm). Wrinkled elytra, shining green infused with red. Back of prothorax has two large black spots. Freshly emerged specimens covered with yellow bloom.
BEHAVIOR
Sits on leaves in open areas and along roadsides; rolls leaves.
FEEDING ECOLOGY AND DIET
Feeds on the leaves of Dichaetanthera cordifolia, a small tree in the family Melastomataceae.
REPRODUCTIVE BIOLOGY
Females lay their eggs on leaves; the leaves are rolled up into a tube to protect and nourish the larvae.
CONSERVATION STATUS
Not threatened.
Scarabaeus sacer
Lytta vesicatoria
Trachelophorus giraffa
Alaus oculatus
Ulochaetes leoninus
Euchroma gigantea
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Order: Coleoptera
Priacma serrata
Zopherus chilensis
Dineutus discolor
DISTRIBUTION
Mexico to Argentina and the Antilles.
HABITAT
Tropical forests.
BEHAVIOR
Common on trunks of living or dead bombacaceous trees.
FEEDING ECOLOGY AND DIET
Larvae bore through trunks of dead bombacaceous trees.
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
Elytra are made into jewelry and ornaments by peoples in Central and South America; adults are eaten by Tzeltal-Mayan Indians in Chiapas, Mexico.
Titanic longhorn beetle
Titanus gigantea
FAMILY
Cerambycidae
TAXONOMY
Titanus gigantea Linnaeus, 1758, “Cayania.”
OTHER COMMON NAMES
None known.
PHYSICAL CHARACTERISTICS
Grows up to 6.7 in (17 cm). Dark brown to black with faint, powerful mandibles and longitudinal ridges on the elytra.
Vol. 3: Insects
DISTRIBUTION
Colombia, Ecuador, Peru, French Guiana, and northern Brazil.
HABITAT
Tropical forests.
BEHAVIOR
Adults are attracted to lights at night.
FEEDING ECOLOGY AND DIET
Larvae probably feed in rotten wood.
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
One of the world’s largest beetles, specimens are highly prized and sell for hundreds of dollars.
Lion beetle
Ulochaetes leoninus
FAMILY
Cerambycidae
TAXONOMY
Ulochaetes leoninus LeConte, 1854, “Prairie Pass, Oregon Territory.”
OTHER COMMON NAMES
None known.
PHYSICAL CHARACTERISTICS
Grows to 0.67–0.98 in (17–25 mm); hairy with black and yellow markings and short elytra.
DISTRIBUTION
Pacific coast, from British Columbia to southern California.
HABITAT
Pine forests.
BEHAVIOR
Look, sound, and behave like bumble bees.
FEEDING ECOLOGY AND DIET
Larvae bore into sapwood of conifers.
REPRODUCTIVE BIOLOGY
Eggs are laid at the base of standing dead trees and stumps.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
Interesting example of physical and behavioral mimicry.
Cupedid beetle
Priacma serrata
FAMILY
Cupedidae
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Vol. 3: Insects
TAXONOMY
Cupes serrata LeConte, 1861, “East of Fort Colville,” Oregon.
OTHER COMMON NAMES
English: cupesid beetle.
PHYSICAL CHARACTERISTICS
Reaches 0.4–0.9 in (11–22 mm). Irregularly marked with grayishblack scales; elytra with square pits.
DISTRIBUTION
Western North America.
HABITAT
Under tree bark in coniferous forests.
BEHAVIOR
During daylight hours, males sometimes are attracted to sheets freshly laundered with bleach.
FEEDING ECOLOGY AND DIET
Probably feed on fungus in rotten wood.
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
A primitive family of beetles similar in appearance to the extinct Tshekardoleidae.
Great water beetle
Dytiscus marginalis
FAMILY
Dytiscidae
TAXONOMY
Dytiscus marginalis Linné, 1758, “Europae.”
OTHER COMMON NAMES
English: Great water diving beetle.
PHYSICAL CHARACTERISTICS
Grows up to 1.4 in (35 mm). Prothorax and elytra have pale borders; male’s elytra is smooth and female’s usually grooved.
DISTRIBUTION
Europe.
HABITAT
Lakes and ponds with muddy bottoms.
BEHAVIOR
Obtains oxygen by breaking the water surface with the tip of the abdomen and trapping a bubble under the elytra.
FEEDING ECOLOGY AND DIET
Preys on aquatic insects, mollusks, crustaceans, and even tadpoles and small fish.
Order: Coleoptera
Lucanus cer vus
Dytiscus marginalis
REPRODUCTIVE BIOLOGY
Eggs are deposited singly on stems of aquatic plants. Larvae molt three times in 35–40 days; pupation takes place in damp ground next to water. One generation per year
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
One of the largest and most intensely studied European water beetles.
Eyed click beetle
Alaus oculatus
FAMILY
Elateridae
TAXONOMY
Elater oculatus Linné, 1758, “America septentrionalis.”
OTHER COMMON NAMES
None known.
PHYSICAL CHARACTERISTICS
Grows to 0.98–1.96 in (25–50 mm). Pronotum with two black spots encircled with white scales.
DISTRIBUTION
Eastern North America.
HABITAT
Woodlands.
BEHAVIOR
Adults found on hardwoods or under bark.
FEEDING ECOLOGY AND DIET
Larvae live in rotten hardwood. Adults and larvae feed on the larvae of wood-boring beetles.
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Order: Coleoptera
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
None known.
Whirligig beetle
Dineutus discolor
FAMILY
Gyrinidae
TAXONOMY
Dineutus discolor Aube, 1838, “États-Unis d’Amerique.”
OTHER COMMON NAMES
English: Gyrinids, apple bugs.
PHYSICAL CHARACTERISTICS
Reaches 0.4–0.5 in (11–13 mm). Black and shining, with pale underside.
DISTRIBUTION
Eastern North America and Mexico.
HABITAT
Surface of slow-moving ponds and streams.
BEHAVIOR
Lives singly or in groups.
FEEDING ECOLOGY AND DIET
Preys on insects trapped on the water surface.
REPRODUCTIVE BIOLOGY
Eggs are laid on submerged plants. Aquatic larvae prey on small invertebrates. Pupate in moist soil near water. Adults overwinter in debris at the edge of water.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
None known.
Pink glowworm
Microphotus angustus
FAMILY
Lampyridae
TAXONOMY
Microphotus angustus LeConte, 1874, “Mariposa, California.”
OTHER COMMON NAMES
None known.
PHYSICAL CHARACTERISTICS
Males reach 0.2 in (6 mm) and females 0.4–0.5 in (11–13 mm). Pink females are larviform; males look like fireflies with large eyes.
Vol. 3: Insects
Dynastes hercules
Microphotus angustus
DISTRIBUTION
California and Oregon.
HABITAT
Forests and moist canyons in chaparral.
BEHAVIOR
Females hang with their heads pointed upward on rocks and stumps and “call” with continuous light. Males seldom are seen and flash weakly only when disturbed.
FEEDING ECOLOGY AND DIET
Unknown; larvae probably prey on small invertebrates.
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
None known.
European stag beetle
Lucanus cervus
FAMILY
Lucanidae
TAXONOMY
Scarabaeus cervus Linnaeus, 1758, “Europae.”
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Vol. 3: Insects
OTHER COMMON NAMES
German: Donnerkafer, Hausbrenner, Feueranzunder, Köhler,
Feuerschröter.
PHYSICAL CHARACTERISTICS
Males reach 1.4–2.95 in (35–75 mm) and females 1.2–1.8 in (30–45 mm). Dark brownish–black beetle. The male has a broad head and antler-like mandibles; female is smaller and more stout, with relatively small mandibles.
DISTRIBUTION
Central, southern, and western Europe; Asia Minor; Syria.
HABITAT
Old oak forests.
BEHAVIOR
Males use mandibles against rival males over females.
FEEDING ECOLOGY AND DIET
Adults feed on sap; larvae eat rotting wood.
REPRODUCTIVE BIOLOGY
Eggs are laid in old wood, larva take three to five years to mature. Adult matures in autumn but overwinters in pupal case.
CONSERVATION STATUS
Collection of this species is forbidden in several European countries.
SIGNIFICANCE TO HUMANS
Historically a symbol of evil and bad luck.
Spanish fly
Lytta vesicatoria
FAMILY
Meloidae
TAXONOMY
Meloe vesicatoria Linné, 1758. Type locality not specified.
OTHER COMMON NAMES
None known.
PHYSICAL CHARACTERISTICS
Reaches 0.5–0.9 in (12–22 mm); slender, soft-bodied metallic golden-green beetle.
DISTRIBUTION
Throughout southern Europe and eastward to Central Asia and Siberia.
HABITAT
Scrublands and woods.
BEHAVIOR
When disturbed, meloids release the blistering agent cantharidin from their leg joints.
FEEDING ECOLOGY AND DIET
Adults feed on leaves of ash, lilac, amur privet, and white willow trees; larvae are parasitic on the brood of ground nesting bees.
REPRODUCTIVE BIOLOGY
Develop by hypermetamorphosis, a type of complete metamorphosis in which first larval instar (the triungulin) is very active,
Order: Coleoptera
while remaining instars are more sedentary and grublike. Eggs are laid near the entrance of host bee’s nest; triungulins crawl into nest on their own.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
Bodies are filled with the toxin cantharidin; elytra were once pulverized and marketed as an aphrodisiac as well as a cure for various ailments.
Hercules beetle
Dynastes hercules
FAMILY
Scarabaeidae
TAXONOMY
Scarabaeus hercules Linné, 1758, “America.”
OTHER COMMON NAMES
French: Scieurs de long; Spanish: Tijeras.
PHYSICAL CHARACTERISTICS
Males reach 5.9–6.7 in (15–17 cm), up to half of which corresponds to the thoracic horn.
DISTRIBUTION
Mexico, Central and northern South America, Guadeloupe, and the Dominican Republic.
HABITAT
Humid tropical forests.
BEHAVIOR
Adults are attracted to oozing sap and sweet fruits; larvae develop in rotten logs.
FEEDING ECOLOGY AND DIET
Nocturnal and frequently attracted to lights.
REPRODUCTIVE BIOLOGY
Males defend feeding sights that will attract females; horns of males are used to grapple with other males over females.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
Ingesting the horn is thought by some people to increase their sexual potency.
Sacred scarab
Scarabaeus sacer
FAMILY
Scarabaeidae
TAXONOMY
Scarabaeus sacer Linné, 1758, “Aegypto.”
OTHER COMMON NAMES
None known.
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Order: Coleoptera
PHYSICAL CHARACTERISTICS
Grows to 0.98–1.2 in (25–30 mm). Black, with rakelike head and forelegs.
DISTRIBUTION
Mediterranean region and central Europe.
HABITAT
Steppe, forest-steppe, and semi-desert.
BEHAVIOR
Adults track dung by smell as food for themselves and their offspring. The female stands head down and rolls a dung ball with the second and third pairs of legs; ball is buried as food for larva.
FEEDING ECOLOGY AND DIET
Adults use membranous mandibles to strain fluids, molds, and other suspended particles as food from dung; larvae eat solid dung.
REPRODUCTIVE BIOLOGY
Single humpbacked larva feeds and pupates inside buried dung ball.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
Symbol of the ancient Egyptian sun god Ra. Scarab jewelry is worn today as a good luck charm. Significant recyclers of animal waste.
Vol. 3: Insects
American burying beetle
Nicrophorus americanus
FAMILY
Silphidae
TAXONOMY
Nicrophorus americanus Olivier, 1790, “Amérique septentrionale.”
OTHER COMMON NAMES
None known.
PHYSICAL CHARACTERISTICS
Reaches 0.8–1.4 in (20–35 mm). Black, with orange antennal club. Head and pronotum have central orange spot; and elytra have four wide spots.
DISTRIBUTION
Formerly throughout eastern North America; restricted now to isolated populations in the Midwest, with populations reintroduced to Rhode Island and Massachusetts.
HABITAT
Woodlands, grassland prairies, forest edge, and scrubland.
BEHAVIOR
Provides parental care for its young.
FEEDING ECOLOGY AND DIET
Scavenges and buries carrion.
REPRODUCTIVE BIOLOGY
Mating pair prepares carrion as food for themselves and their larvae.
Ocypus olens
Nicrophorus americanus Titanus gigantea
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Vol. 3: Insects
CONSERVATION STATUS
Listed as Endangered by IUCN and by the U.S. Fish and Wildlife Service.
SIGNIFICANCE TO HUMANS
Symbolic of habitat destruction and modification throughout eastern North America.
Devil’s coach-horse
Ocypus olens
FAMILY
Staphylinidae
TAXONOMY
Staphylinus olens Müller, 1764. Type locality not specified.
OTHER COMMON NAMES
English: Rove beetle.
PHYSICAL CHARACTERISTICS
Grows to 0.9–1.3 in (22–33 mm). Black, with short elytra exposing abdominal segments.
DISTRIBUTION
Lower elevations of Europe, Russia, Turkey, North Africa, and the Canary Islands; established in parts of North America.
HABITAT
Under stones, damp leaves, and moss.
BEHAVIOR
When alarmed, the beetle spreads its powerful jaws and curls its abdomen over its back to emit a foul-smelling brown fluid.
FEEDING ECOLOGY AND DIET
Adults and larvae prey on soil-dwelling invertebrates.
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
Order: Coleoptera
SIGNIFICANCE TO HUMANS
Once a symbol of evil associated with death.
Ma’kech
Zopherus chilensis
FAMILY
Zopheridae
TAXONOMY
Zophorus [sic] chilensis Gray, 1832. Type locality not specified.
OTHER COMMON NAMES
English: Jeweled beetle.
PHYSICAL CHARACTERISTICS
Grows to 1.3–1.6 in (34–40 mm). Extremely hard exoskeleton; rough back is white with irregular black blotches.
DISTRIBUTION
Southern Mexico to Venezuela and Colombia.
HABITAT
Found on bark of dead trees.
BEHAVIOR
When disturbed, adults tuck in their legs and play dead.
FEEDING ECOLOGY AND DIET
Larvae presumably feed on fungal hyphae in rotten wood; adults eat cereals in captivity.
REPRODUCTIVE BIOLOGY
Nothing is known.
CONSERVATION STATUS
Not threatened.
SIGNIFICANCE TO HUMANS
Living beetles are adorned with beads, tethered with a gold chain, and worn as living costume jewelry as traditional reminder of an ancient Yucatecan legend.
Resources
Books
Arnett, Ross H., Jr., and Michael C. Thomas, eds. American Beetles. Vol. 1, Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. Boca Raton, FL: CRC Press, 2001.
Arnett, Ross H., Jr., Michael C. Thomas, Paul E. Skelley, and J. Howard Frank, eds. American Beetles. Vol. 2, Polyphaga: Scarabaeoidea through Curculionidae. Boca Raton, FL: CRC Press. 2002.
Crawford, C. S. Biology of Desert Invertebrates. Berlin: SpringerVerlag, 1981.
Crowson, R. A. The Biology of the Coleoptera. London: Academic Press, 1981.
Elias, Scott A. Quaternary Insects and Their Environments.
Washington, DC: Smithsonian Institution Press, 1994.
Evans, Arthur V., and Charles L. Bellamy. An Inordinate Fondness for Beetles. Berkeley: University of California Press, 2000.
Evans, Arthur V., and J. N. Hogue. Introduction to California Beetles. Berkeley: University of California Press, in press.
Evans, D. L., and J. O. Schmidt, eds. Insect Defenses: Adaptive Mechanisms and Strategies of Prey and Predators. Albany: State University of New York Press, 1990.
Klausnitzer, B. Beetles. New York: Exeter Books, 1983.
Lawrence, J. F., and E. B. Britton. Australian Beetles. Carlton, Australia: Melbourne University Press, 1994.
Lawrence, J. F., and A. F. Newton, Jr. “Families and Subfamilies of Coleoptera (with Selected Genera, Notes, References and Data on Family-Group Names).” In Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating
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Order: Coleoptera
Resources
the 80th Birthday of Roy A. Crowson, edited by J. Pakaluk and S. A. Slipinski. Warsaw, Poland: Muzeum i Instytut Zoologii PAN, 1995.
Meads, M. Forgotten Fauna: The Rare, Endangered, and Protected Invertebrates of New Zealand. Wellington, New Zealand: Department of Scientific and Industrial Research, 1990.
Pakaluk, J., and S. A. Slipinski, eds. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson. Warsaw, Poland: Muzeum i Instytut Zoologii PAN, 1995.
Stehr, Frederick W., ed. Immature Insects. Vol. 2. Dubuque, IA: Kendall/Hunt Publishing Company, 1991.
White, R. E. A Field Guide to the Beetles of North America.
Boston: Houghton Mifflin, 1983.
Periodicals
Alcock, J. “Postinsemination Associations between Males and Females in Insects: The Mate-Guarding Hypothesis.”
Annual Review of Entomology 39 (1994): 1–21.
Beutel, R. “Über Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga.” Abhandlungen des Naturwissenshaftlichen Vereins in Hamburg 31 (1997): 1–164.
Beutel, R., and F. Haas. “Phylogenetic Relationships of the Suborders of Coleoptera (Insecta).” Cladistics 16 (2000): 103–141.
Caterino, M. S., V. L. Shull, P. M. Hammond, and A. P. Vogler. “Basal Relationships of Coleoptera Inferred from 18S rDNA Sequences.” Zoologica Scripta 31 (2002): 41–49.
Eberhard, William G. “Horned Beetles.” Scientific American 242, no. 3 (1980): 166–182.
Emlen, D. J. “Integrating Development with Evolution: A Case Study with Beetle Horns.” Bioscience 50, no. 5 (2000): 403–418.
Farrell, Brian D. “‘Inordinate Fondness’ Explained: Why Are There So Many Beetles?” Science 281 (1998): 555–559.
Vol. 3: Insects
Hadley, N. F. “Beetles Make Their Own Waxy Sunblock.” Natural History 102, no. 8 (1993): 44–45.
Lomolino, Mark V., J. C. Creighton, G. D. Schnell, and D. L. Certain. “Ecology and Conservation of the Endangered American Burying Beetle (Nicrophorus americanus).” Conservation Biology 9, no. 3 (1995): 605–614.
McIver, J. D., and G. Stonedahl. “Myrmecomorphy: Morphological and Behavioral Mimicry of Ants.” Annual Review of Entomology 38 (1993): 351–379.
Milne, L. J., and M. J. Milne. “The Social Behavior of Burying Beetles.” Scientific American 235, no. 2 (1976): 84–89.
Murlis, J. “Odor Plumes and How Insects Use Them.” Annual Review of Entomology 37 (1992): 505–532.
Rettenmeyer, C. W. “Insect Mimicry.” Annual Review of Entomology 15 (1970): 43–74.
Other
“The Balfour-Browne Club.” October 1996 [April 14, 2003].
<http://www.lifesci.utexas.edu/faculty/sjasper/beetles/
BBClub.htm>.
“Beetles (Coleoptera) and Coleopterists.” [April 14, 2003].
<http://www.zin.ru/Animalia/Coleoptera/eng>.
“Coleoptera Home Page.” March 28, 2003 [April 14, 2003].
<http://www.coleoptera.org>.
“The Coleopterist.” [April 14, 2003]. <http://www.coleopterist
.org.uk>.
“Coleopterists Society.” [April 14, 2003]. <http://www
.coleopsoc.org>.
“The Japan Coleopterological Society.” [April 14, 2003]. <http://www.mus-nh.city.osaka.jp/shiyake/j-coleopt-soc
.html>.
“Wiener Coleopterologen Verein (Vienna Coleopterists Society).” September 7, 2001 [April 14, 2003]. <http://www
.nhm-wien.ac.at/NHM/2Zoo/coleoptera/wcv_e.html>.
Arthur V. Evans, DSc
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