Etymology[edit]

Coleoptera comes from the Greek koleopteros, literally "sheath-wing", from koleos meaning
"sheath", and pteron, meaning "wing". The name was given to the group byAristotle for
their elytra, hardened shield-like forewings. The English name "beetle" comes from the Old
English word bitela, literally meaning small biter, deriving from the word bitel, which means
biting. This word is related to the word btan (to bite)[2][3] The name also derives from
the Middle English word betylle from Old Englishbitula (also meaning to bite).[4] Another Old
English name for beetle is ceafor, chafer, used in names such as cockchafer, from the ProtoGermanic *kabraz- (compare German Kfer).[5] These terms have been in use since the 12th
century.[4] In addition to names including the words "beetle" or "chafer", many groups of
Coleoptera have common names such as fireflies, June bugs, ladybugs and weevils.[6]

Taxonomy[edit]

A museum display, showing a little of the diversity of beetles

The Coleopterans include more species than any other order, constituting nearly 25% of all
known types of animal life forms.[6][7][8] About 450,000 species of beetles occur
representing about 40% of all known insects.[9] Such a large number of species poses special
problems for classification, with some families consisting of thousands of species and needing
further division into subfamilies and tribes. This immense number of species allegedly led
evolutionary biologist J. B. S. Haldane to quip, when some theologians asked him what could
be inferred about the mind of the Creator from the works of His Creation, that God displayed
"an inordinate fondness for beetles".[10]
Polyphaga is the largest suborder, containing more than 300,000 described species in more
than 170 families, including rove beetles (Staphylinidae), scarab beetles
(Scarabaeidae), blister beetles (Meloidae), stag beetles (Lucanidae) and true weevils
(Curculionidae).[8][11] These beetles can be identified by the presence of
cervical sclerites (hardened parts of the head used as points of attachment for muscles)
absent in the other suborders.[citation needed] The suborder Adephaga contains about 10
families of largely predatory beetles, includes ground
beetles (Carabidae), Dytiscidae and whirligig beetles (Gyrinidae). In these beetles,

the testes are tubular and the first abdominal sternum (a plate of the exoskeleton) is divided
by the hind coxae (the basal joints of the beetle's legs).[citation needed] Archostemata contains
four families of mainly wood-eating beetles, includingreticulated beetles (Cupedidae) and
the telephone-pole beetle.[citation needed] Myxophaga contains about 100 described species in
four families, mostly very small, including Hydroscaphidae and the genus Sphaerius.[citation
needed]

Evolution[edit]
The oldest known insect that unequivocally resembles species of Coleoptera date back to
the Lower Permian (270 mya), though it instead has 13-segmented antennae,elytra with more
fully developed venation and more irregular longitudinal ribbing, and an abdomen
and ovipositor extending beyond the apex of the elytra. At the end of the Permian, the
biggest mass extinction in history took place, collectively called the PermianTriassic
extinction event (P-Tr): 30% of all insect species became extinct; however, it is the only mass
extinction of insects in Earth's history until today.[12]
Due to the P-Tr extinction, the fossil record of insects only includes beetles from the Lower
Triassic (220 million years ago). Around this time, during the Late Triassic,mycetophagous, or
fungus-feeding species (e.g. Cupedidae) appear in the fossil record. In the stages of the Upper
Triassic, representatives of the algophagous, or algae-feeding species
(e.g. Triaplidae and Hydrophilidae) begin to appear, as well as predatory water beetles. The
first primitive weevils appear (e.g. Obrienidae), as well as the first representatives of the rove
beetles (e.g. Staphylinidae), which show no marked difference in morphology compared to
recent species.[12]

Baltic amber inclusions from the Eocene, 50 million years ago

During the Jurassic (210 to 145 million years ago), a dramatic increase in the known diversity
of family-level Coleoptera occurred,[12] including the development and growth of carnivorous
and herbivorous species. Species of the superfamilyChrysomeloidea are believed to have
developed around the same time, which include a wide array of plant hosts ranging
fromcycads and conifers, to angiosperms.[13] Close to the Upper Jurassic, the portion of
the Cupedidae decreased, but at the same time the diversity of the early plant-eating, or
phytophagous species increased. Most of the recent phytophagous species of Coleoptera feed
on flowering plants or angiosperms. The increase in diversity of the angiosperms is also
believed to have influenced the diversity of the phytophagous species, which doubled during
the Middle Jurassic. However, doubts have been raised recently, since the increase of the
number of beetle families during the Cretaceous does not correlate with the increase of the
number of angiosperm species.[14] Also around the same time, numerous primitive weevils
(e.g. Curculionoidea) and click beetles (e.g. Elateroidea) appeared. Also, the first jewel beetles
(e.g. Buprestidae) are present, but they were rather rare until the Cretaceous.[15][16][17] The
first scarab beetles appeared around this time, but they were not coprophagous (feeding
upon fecal matter), instead presumably feeding upon the rotting wood with the help of
fungus; they are an early example of a mutualistic relationship.

The Cretaceous included the initiation of the most recent round of southern landmass
fragmentation, via the opening of the southern Atlantic ocean and the isolation of New
Zealand, while South America, Antarctica, and Australia grew more distant. [13] During the
Cretaceous, the diversity of Cupedidae and Archostemata decreased considerably.
Predatory ground beetles (Carabidae) and rove beetles (Staphylinidae) began to distribute
into different patterns; whereas the Carabidae predominantly occurred in the warm regions,
the Staphylinidae and click beetles (Elateridae) preferred many areas with temperate
climates. Likewise, predatory species of Cleroideaand Cucujoidea hunted their prey under the
bark of trees together with the jewel beetles (Buprestidae). The jewel beetles' diversity
increased rapidly during the Cretaceous, as they were the primary consumers of wood,
[18] while longhorn beetles (Cerambycidae) were rather rare, and their diversity increased
only towards the end of the Upper Cretaceous.[12] The first coprophagous beetles have been
recorded from the Upper Cretaceous,[19] and are believed to have lived on the excrement of
herbivorous dinosaurs, but discussion is still ongoing as to whether the beetles were always
tied to mammals during their development.[20] Also, the first species with an adaption of both
larvae and adults to the aquatic lifestyle are found. Whirligig beetles (Gyrinidae) were
moderately diverse, although other early beetles (e.g. Dytiscidae) were less, with the most
widespread being the species of Coptoclavidae, which preyed on aquatic fly larvae.[12]
Between the Paleogene and the Neogene is when today's beetles developed. During this time,
the continents began to be located closer to where they are today. Around 5 million years
ago, the land bridge between South America and North America was formed, and the fauna
exchange between Asia and North America started. Though many recent genera and species
already existed during the Miocene, their distribution differed considerably from today's. [12]

Fossil record[edit]

Fossil buprestid beetle from theEocene Messel pit, which retains itsstructural color

A 2007 study based on DNA of living beetles and maps of likely beetle evolution indicated
beetles may have originated during the Lower Permian, up to 285 million years ago.[8] In
2009, a fossil beetle was described from the Pennsylvanian of Mazon Creek, Illinois, pushing
the origin of the beetles to an earlier date, 318 to 299 million years ago.[21] Fossils from this
time have been found in Asia and Europe, for instance in the red slate fossil beds of
Niedermoschel near Mainz, Germany.[22] Further fossils have been found in Obora, Czech
Republic and Tshekarda in the Ural mountains, Russia.[23] However, there are only a few
fossils from North America before the middle Permian, although both Asia and North America
had been united toEuramerica. The first discoveries from North America made in the
Wellington formation of Oklahoma were published in 2005 and 2008.[12][24]
As a consequence of the PermianTriassic extinction event, the fossil record of insects is
scant, including beetles from the Lower Triassic. [25] However, a few exceptions are noted, as
in Eastern Europe; at the Babiy Kamen site in the Kuznetsk Basin, numerous beetle fossils

were discovered, even entire specimen of the infraorders Archostemata (e.g. Ademosynidae,
Schizocoleidae), Adephaga (e.., Triaplidae, Trachypachidae) and Polyphaga (e.g.
Hydrophilidae, Byrrhidae, Elateroidea) and in nearly a perfectly preserved condition.
[26] However, species from the families Cupedidae and Schizophoroidae are not present at this
site, whereas they dominate at other fossil sites from the Lower Triassic. Further records are
known from Khey-Yaga, Russia, in the Korotaikha Basin.[12] There are many important sites
from the Jurassic, with more than 150 important sites with beetle fossils, the majority being
situated in Eastern Europe and North Asia. In North America and especially in South America
and Africa, the number of sites from that time period is smaller, and the sites have not been
exhaustively investigated yet. Outstanding fossil sites include Solnhofen in Upper Bavaria,
Germany,[27] Karatau in South Kazakhstan,[28] the Yixian formation in Liaoning, North China,
[29] as well as the Jiulongshan formation and further fossil sites in Mongolia. In North America
there are only a few sites with fossil records of insects from the Jurassic, namely the shell
limestone deposits in the Hartford basin, the Deerfield basin and the Newark basin. [12][30]