See subgroups of the order Coleoptera
Beetles are a group of insects that form the order Coleoptera, in the
superorder Endopterygota. Their front pair of wings is hardened into
wing-cases, elytra, distinguishing them from most other insects. The
Coleoptera, with about 400,000 species, is the largest of all orders,
constituting almost 40% of described insects and 25% of all known
animal life-forms; new species are discovered frequently. The largest
of all families, the
Curculionidae (weevils) with some 70,000 member
species, belongs to this order. They are found in almost every habitat
except the sea and the polar regions. They interact with their
ecosystems in several ways: beetles often feed on plants and fungi,
break down animal and plant debris, and eat other invertebrates. Some
species are serious agricultural pests, such as the Colorado potato
beetle, while others such as
Coccinellidae (ladybirds or ladybugs) eat
aphids, scale insects, thrips, and other plant-sucking insects that
Beetles typically have a particularly hard exoskeleton including the
elytra, though some such as the rove beetles have very short elytra
while blister beetles have softer elytra. The general anatomy of a
beetle is quite uniform and typical of insects, although there are
several examples of novelty, such as adaptations in water beetles
which trap air bubbles under the elytra for use while diving. Beetles
are endopterygotes, which means that they undergo complete
metamorphosis, with a series of conspicuous and relatively abrupt
changes in body structure between hatching and becoming adult after a
relatively immobile pupal stage. Some, such as stag beetles, have a
marked sexual dimorphism, the males possessing enormously enlarged
mandibles which they use to fight other males. Many beetles are
aposematic, with bright colours and patterns warning of their
toxicity, while others are harmless Batesian mimics of such insects.
Many beetles, including those that live in sandy places, have
Beetles are prominent in human culture, from the sacred scarabs of
ancient Egypt to beetlewing art and use as pets or fighting insects
for entertainment and gambling. Many beetle groups are brightly and
attractively coloured making them objects of collection and decorative
displays. Over 300 species are used as food, mostly as larvae; species
widely consumed include mealworms and rhinoceros beetle larvae.
However, the major impact of beetles on human life is as agricultural,
forestry, and horticultural pests. Serious pests include the boll
weevil of cotton, the Colorado potato beetle, the coconut hispine
beetle, and the mountain pine beetle. Most beetles, however, do not
cause economic damage and many, such as the lady beetles and dung
beetles are beneficial by helping to control insect pests.
2 Distribution and diversity
3.1 Late Paleozoic
5 External morphology
Anatomy and physiology
6.1 Digestive system
6.2 Nervous system
6.3 Respiratory system
6.4 Circulatory system
6.5 Specialized organs
7 Reproduction and development
7.2 Life cycle
Pupa and adult
8.3 Parental care
9.1 Anti-predator adaptations
Mimicry and aposematism
9.1.3 Other defences
9.5 Tolerance of extreme environments
10 Relationship to humans
10.1 In ancient cultures
10.2 As pests
10.3 As beneficial resources
10.4 As food
10.5 As biodiversity indicators
10.6 In art and adornment
10.7 In entertainment
10.8 As pets
10.9 As things to collect
10.10 As inspiration for technologies
10.11 In conservation
13 Further reading
14 External links
Coleoptera at the Staatliches Museum für Naturkunde Karlsruhe,
The name of the taxonomic order, Coleoptera, comes from the Greek
koleopteros (κολεόπτερος), given to the group by Aristotle
for their elytra, hardened shield-like forewings, from koleos, sheath,
and pteron, wing. The English name beetle comes from the Old English
word bitela, little biter, related to bītan (to bite), leading
Middle English betylle. Another
Old English name for beetle is
ceafor, chafer, used in names such as cockchafer, from the
Proto-Germanic *kabraz- (compare German Käfer).
Distribution and diversity
Beetles are by far the largest order of insects: the roughly 400,000
species make up about 40% of all insect species so far described, and
about 25% of all animals. A 2015 study provided
four independent estimates of the total number of beetle species,
giving a mean estimate of some 1.5 million with a "surprisingly narrow
range" spanning all four estimates from a minimum of 0.9 to a
maximum of 2.1 million beetle species. The four estimates made use of
host-specificity relationships (1.5 to 1.9 million), ratios with other
taxa (0.9 to 1.2 million), plant:beetle ratios (1.2 to 1.3), and
extrapolations based on body size by year of description (1.7 to 2.1
Beetles are found in nearly all habitats, including freshwater and
coastal habitats, wherever vegetative foliage is found, from trees and
their bark to flowers, leaves, and underground near roots - even
inside plants in galls, in every plant tissue, including dead or
The heaviest beetle, indeed the heaviest insect stage, is the larva of
the goliath beetle,
Goliathus goliatus, which can attain a mass of at
least 115 g (4.1 oz) and a length of 11.5 cm
(4.5 in). Adult male goliath beetles are the heaviest beetle in
its adult stage, weighing 70–100 g (2.5–3.5 oz) and
measuring up to 11 cm (4.3 in). Adult elephant beetles,
Megasoma elephas and
Megasoma actaeon often reach 50 g
(1.8 oz) and 10 cm (3.9 in).
The longest beetle is the
Hercules beetle Dynastes hercules, with a
maximum overall length of at least 16.7 cm (6.6 in)
including the very long pronotal horn. The smallest recorded beetle
and the smallest free-living insect (as of 2015), is the featherwing
Scydosella musawasensis which may measure as little as
325 µm in length.
Titan beetle, Titanus giganteus, a tropical longhorn, is one of the
largest and heaviest insects in the world.
Scydosella musawasensis, the smallest known beetle: scale bar (right)
is 50 μm.
Hercules beetle, Dynastes hercules ecuatorianus, the longest of all
The oldest known fossil insect that unequivocally resembles a
Coleopteran is from the Lower
Permian Period about 270 million
years ago (mya), though these members of the family Tshekardocoleidae
have 13-segmented antennae, elytra with more fully developed venation
and more irregular longitudinal ribbing, and abdomen and ovipositor
extending beyond the apex of the elytra. In the Permian–Triassic
extinction event at the end of the Permian, some 30% of all insect
species became extinct, so the fossil record of insects only includes
beetles from the Lower
Triassic 220 mya. Around this time, during the
Late Triassic, fungus-feeding species such as
Cupedidae appear in the
fossil record. In the stages of the Upper Triassic, alga-feeding
insects such as Triaplidae and
Hydrophilidae begin to appear,
alongside predatory water beetles. The first weevils, including the
Obrienidae, appear alongside the first rove beetles (Staphylinidae),
which closely resemble recent species. Some entomologists are
sceptical that such early insects are so closely related to
present-day species, arguing that this is extremely unlikely; for
example, the structure of the metepisternum suggests that the
Obrienidae could be Archostemata, not weevils at all, despite fossils
with weevil-like snouts.
Beetle genera were mainly saprophages in the
Permian and Triassic.
During the Jurassic, herbivorous and then carnivorous genera became
more common. In the Cenozoic, genera at all three trophic levels
became far more numerous.
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 mya. Fossils from this time have been found
in Asia and Europe, for instance in the red slate fossil beds of
Niedermoschel near Mainz, Germany. Further fossils have been found
in Obora, Czech Republic and Tshekarda in the Ural mountains,
Russia. However, there are only a few fossils from North America
before the middle Permian, although both Asia and North America had
been united to Euramerica. The first discoveries from North America
made in the Wellington formation of Oklahoma were published in 2005
As a consequence of the Permian–
Triassic extinction event, the
fossil record of insects is scant, including beetles from the Lower
Triassic. However, there are a few exceptions, such as in Eastern
Europe. At the Babiy Kamen site in the Kuznetsk Basin, numerous beetle
fossils were discovered, including entire specimens of the infraorders
Archostemata (e.g. Ademosynidae, Schizocoleidae),
Triaplidae, Trachypachidae) and
Polyphaga (e.g. Hydrophilidae,
Byrrhidae, Elateroidea). 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 such as
Khey-Yaga, Russia, in the Korotaikha Basin.
Jurassic (210 to 145 mya), there was a dramatic
increase in the diversity of beetle families, including the
development and growth of carnivorous and herbivorous species. The
Chrysomeloidea diversified around the same time, feeding on a wide
array of plant hosts from cycads and conifers to angiosperms.
Close to the Upper Jurassic, the
Cupedidae decreased, but the
diversity of the early plant-eating species increased. Most recent
plant-eating beetles feed on flowering plants or angiosperms, whose
success contributed to a doubling of plant-eating species during the
Middle Jurassic. However, the increase of the number of beetle
families during the
Cretaceous does not correlate with the increase of
the number of angiosperm species. Around the same time, numerous
primitive weevils (e.g. Curculionoidea) and click beetles (e.g.
Elateroidea) appeared. The first jewel beetles (e.g. Buprestidae) are
present, but they remained rare until the Cretaceous. The
first scarab beetles were not coprophagous but presumably fed on
rotting wood with the help of fungus; they are an early example of a
There are more than 150 important fossil sites from the Jurassic, the
majority in Eastern Europe and North Asia. Outstanding sites include
Solnhofen in Upper Bavaria, Germany, Karatau in South
Kazakhstan, the Yixian formation in Liaoning, North China, 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
Cretaceous saw the fragmenting of the southern landmass, with the
opening of the southern Atlantic Ocean and the isolation of New
Zealand, while South America, Antarctica, and
Australia grew more
distant. The diversity of
considerably. Predatory ground beetles (Carabidae) and rove beetles
(Staphylinidae) began to distribute into different patterns; the
Carabidae predominantly occurred in the warm regions, while the
Staphylinidae and click beetles (Elateridae) preferred temperate
climates. Likewise, predatory species of
Cleroidea and Cucujoidea
hunted their prey under the bark of trees together with the jewel
beetles (Buprestidae). The diversity of jewel beetles increased
rapidly, as they were the primary consumers of wood, while
longhorn beetles (Cerambycidae) were rather rare: their diversity
increased only towards the end of the Upper Cretaceous. The first
coprophagous beetles are from the Upper Cretaceous and may have
lived on the excrement of herbivorous dinosaurs. The first species
where both larvae and adults are adapted to an 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
Many fossil sites worldwide contain beetles from the Cretaceous. Most
are in Europe and Asia and belong to the temperate climate zone during
the Cretaceous. Lower
Cretaceous sites include the Crato fossil
beds in the Araripe basin in the Ceará, North Brazil, as well as
overlying Santana formation; the latter was near the equator at that
time. In Spain, important sites are near Montsec and Las Hoyas. In
Australia, the Koonwarra fossil beds of the Korumburra group, South
Gippsland, Victoria, are noteworthy. Major sites from the Upper
Cretaceous include Kzyl-Dzhar in South
Kazakhstan and Arkagala in
Fossil buprestid beetle from the
Eocene (50 mya) Messel pit, which
retains its structural color
Beetle fossils are abundant in the Cenozoic; by the
Quaternary (up to
1.6 mya), fossil species are identical to living ones, while from the
Late Miocene (5.7 mya) the fossils are still so close to modern forms
that they are most likely the ancestors of living species. The large
oscillations in climate during the
Quaternary caused beetles to change
their geographic distributions so much that current location gives
little clue to the biogeographical history of a species. It is evident
that geographic isolation of populations must often have been broken
as insects moved under the influence of changing climate, causing
mixing of gene pools, rapid evolution, and extinctions, especially in
The very large number of beetle species poses special problems for
classification. Some families contain tens of thousands of species,
and need to be divided into subfamilies and tribes. This immense
number led the evolutionary biologist
J. B. S. Haldane
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, "An inordinate fondness
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). These polyphagan beetle groups 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
Adephaga contains about 10 families of largely
predatory beetles, includes ground beetles (Carabidae), water beetles
(Dytiscidae) and whirligig beetles (Gyrinidae). In these insects, 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
Archostemata contains four families of mainly
wood-eating beetles, including reticulated beetles (Cupedidae) and the
telephone-pole beetle. The
Archostemata have an exposed plate
called the metatrochantin in front of the basal segment or coxa of the
Myxophaga contains about 65 described species in four
families, mostly very small, including
Hydroscaphidae and the genus
Sphaerius. The myxophagan beetles are small and mostly
alga-feeders. Their mouthparts are characteristic in lacking galeae
and having a mobile tooth on their left mandible.
The consistency of beetle morphology, in particular their possession
of elytra, has long suggested that Coleoptera is monophyletic, though
there have been doubts about the arrangement of the suborders, namely
the Adephaga, Archostemata,
Polyphaga within that
clade. The twisted-wing parasites, Strepsiptera,
are thought to be a sister group to the beetles, having split from
them in the Early Permian.
Molecular phylogenetic analysis confirms that the Coleoptera are
monophyletic. Duane McKenna et al. (2015) used eight nuclear genes for
367 species from 172 of 183 Coleopteran families. They split the
Adephaga into 2 clades, Hydradephaga and Geadephaga, broke up the
Cucujoidea into 3 clades, and placed the
Lymexyloidea within the
Polyphaga appear to date from the Triassic. Most
extant beetle families appear to have arisen in the Cretaceous.
The cladogram is based on McKenna (2015). The number of species in
each group (mainly superfamilies) is shown in parentheses, and
boldface if over 10,000. English names are given where possible. Dates
of origin of major groups are shown in italics in millions of years
Archostemata 160mya (40)
Myxophaga 220mya (94)
Hydradephaga (5560) e.g.
Dytiscidae (diving beetles)
Geadephaga (35000) e.g.
Carabidae (ground beetles)
Derodontoidea 200mya (800)
Staphylinidae 195mya (48000, rove beetles)
Scarabaeoidea 145mya (35000, scarabs, stag beetles, etc)
Hydrophiloidea (2800, water scavenger beetles)
Histeroidea (3800, clown beetles)
Buprestoidea (3000, jewel beetles)
Byrrhoidea (400, pill and turtle beetles, etc)
Elateroidea (23000, click and soldier beetles, fireflies)
Bostrichoidea (3000, deathwatch, powderpost and skin beetles)
Coccinelloidea (6000, ladybirds or lady beetles)
Tenebrionoidea 180mya (35000, leaf/flower beetles, etc) and
Cleroidea (9900, checkered beetles and allies)
Chrysomelidae (35000, leaf beetles)
Cerambycidae (25000, longhorn beetles)
Curculionoidea (70000, weevils)
Beetle body structure, using cockchafer. A: head, B: thorax, C:
abdomen. 1: antenna, 2: compound eye, 3: femur, 4: elytron (wing
cover), 5: tibia, 6: tarsus, 7: claws, 8: mouthparts, 9: prothorax,
10: mesothorax, 11: metathorax, 12: abdominal sternites, 13: pygidium.
Beetles are generally characterized by a particularly hard exoskeleton
and hard forewings (elytra) not usable for flying. Almost all beetles
have mandibles that move in a horizontal plane. The mouthparts are
rarely suctorial, though they are sometimes reduced; the maxillae
always bear palps. The antennae usually have 11 or fewer segments,
except in some groups like the
Cerambycidae (longhorn beetles) and the
Rhipiceridae (cicada parasite beetles). The coxae of the legs are
usually located recessed within a coxal cavity. The genitalic
structures are telescoped into the last abdominal segment in all
Beetle larvae can often be confused with those of
other endopterygote groups. The beetle's exoskeleton is made up of
numerous plates, called sclerites, separated by thin sutures. This
design provides armored defenses while maintaining flexibility. The
general anatomy of a beetle is quite uniform, although specific organs
and appendages vary greatly in appearance and function between the
many families in the order. Like all insects, beetles' bodies are
divided into three sections: the head, the thorax, and the abdomen.
Because there are so many species, identification is quite difficult,
and relies on attributes including the shape of the antennae, the
tarsal formulae[a] and shapes of these small segments on the legs, the
mouthparts, and the ventral plates (sterna, pleura, coxae). In many
species accurate identification can only be made by examination of the
unique male genitalic structures.
The head, having mouthparts projecting forward or sometimes
downturned, is usually heavily sclerotized and is sometimes very
large. The eyes are compound and may display remarkable
adaptability, as in the case of the aquatic whirligig beetles
(Gyrinidae), where they are split to allow a view both above and below
the waterline. Longhorn beetles (Cerambycidae) and weevils have
divided eyes, while many have eyes that are notched, and a few have
ocelli, small, simple eyes usually farther back on the head (on the
vertex); these are more common in larvae than in adults. Ocelli
are found in the adult carpet beetle (Dermestidae), some rove beetles
(Omaliinae), and the Derodontidae.
Polyphylla fullo has distinctive fan-like antennae, one of several
distinct forms for the appendages among beetles.
Beetle antennae are primarily organs of sensory perception and can
detect motion, odour and chemical substances, but may also be used
to physically feel a beetle's environment.
Beetle families may use
antennae in different ways. For example, when moving quickly, tiger
beetles may not be able to see very well and instead hold their
antennae rigidly in front of them in order to avoid obstacles.
Cerambycidae use antennae to balance, and blister beetles may
use them for grasping. Some aquatic beetle species may use antennae
for gathering air and passing it under the body whilst submerged.
Equally, some families use antennae during mating, and a few species
use them for defence. In the cerambycid Onychocerus albitarsis, the
antennae have venom injecting structures used in defence. Antennae
vary greatly in form, sometimes between the sexes, but are often
similar within any given family. Antennae may be clubbed, threadlike,
angled, shaped like a string of beads, comb-like (either on one side
or both, bipectinate), or toothed. The physical variation of antennae
is important for the identification of many beetle groups. The
Curculionidae have elbowed or geniculate antennae. Feather like
flabellate antennae are a restricted form found in the Rhipiceridae
and a few other families. The
Silphidae have a capitate antennae with
a spherical head at the tip. The
Scarabaeidae typically have lamellate
antennae with the terminal segments extended into long flat structures
stacked together. The
Carabidae typically have thread-like antennae.
The antennae arises between the eye and the mandibles and in the
Tenebrionidae, the antennae rise in front of a notch that breaks the
usually circular outline of the compound eye. They are segmented and
usually consist of 11 parts, the first part is called the scape and
the second part is the pedicel. The other segments are jointly called
Beetles have mouthparts like those of grasshoppers. The mandibles
appear as large pincers on the front of some beetles. The mandibles
are a pair of hard, often tooth-like structures that move horizontally
to grasp, crush, or cut food or enemies (see defence, below). Two
pairs of finger-like appendages, the maxillary and labial palpi, are
found around the mouth in most beetles, serving to move food into the
mouth. In many species, the mandibles are sexually dimorphic, with
those of the males enlarged enormously compared with those of females
of the same species.
The thorax is segmented into the two discernible parts, the pro- and
pterothorax. The pterothorax is the fused meso- and metathorax, which
are commonly separated in other insect species, although flexibly
articulate from the prothorax. When viewed from below, the thorax is
that part from which all three pairs of legs and both pairs of wings
arise. The abdomen is everything posterior to the thorax. When
viewed from above, most beetles appear to have three clear sections,
but this is deceptive: on the beetle's upper surface, the middle
section is a hard plate called the pronotum, which is only the front
part of the thorax; the back part of the thorax is concealed by the
beetle's wings. This further segmentation is usually best seen on the
Acilius sulcatus, a diving beetle with hind legs adapted as swimming
The multisegmented legs end in two to five small segments called
tarsi. Like many other insect orders, beetles have claws, usually one
pair, on the end of the last tarsal segment of each leg. While most
beetles use their legs for walking, legs have been variously adapted
for other uses. Aquatic beetles including the
beetles), Haliplidae, and many species of Hydrophilidae, the legs,
often the last pair, are modified for swimming, typically with rows of
long hairs. Male diving beetles have suctorial cups on their forelegs
that they use to grasp females. Other beetles have fossorial legs
widened and often spined for digging. Species with such adaptations
are found among the scarabs, ground beetles, and clown beetles
(Histeridae). The hind legs of some beetles, such as flea beetles
(within Chrysomelidae) and flea weevils (within Curculionidae), have
enlarged femurs that help them leap.
Trichodes alvearius taking off, showing the hard elytra
(forewings adapted as wing-cases) held stiffly away from the flight
The forewings of beetles are not used for flight, but form elytra
which cover the hind part of the body and protect the hindwings. The
elytra are usually hard shell-like structures which must be raised to
allow the hind wings to move for flight. However, in the soldier
beetles (Cantharidae), the elytra are soft, earning this family the
name of leatherwings. Other soft wing beetles include the
net-winged beetle Calopteron discrepans, which has brittle wings that
rupture easily in order to release chemicals for defence.
Beetles' flight wings are crossed with veins and are folded after
landing, often along these veins, and stored below the elytra. A fold
(jugum) of the membrane at the base of each wing is
characteristic. Some beetles have lost the ability to fly. These
include some ground beetles (Carabidae) and some true weevils
(Curculionidae), as well as desert- and cave-dwelling species of other
families. Many have the two elytra fused together, forming a solid
shield over the abdomen. In a few families, both the ability to fly
and the elytra have been lost, as in the glow-worms (Phengodidae),
where the females resemble larvae throughout their lives. The
presence of elytra and wings does not always indicate that the beetle
will fly. For example, the tansy beetle walks between habitats despite
being physically capable of flight.
The abdomen is the section behind the metathorax, made up of a series
of rings, each with a hole for breathing and respiration, called a
spiracle, composing three different segmented sclerites: the tergum,
pleura, and the sternum. The tergum in almost all species is
membranous, or usually soft and concealed by the wings and elytra when
not in flight. The pleura are usually small or hidden in some species,
with each pleuron having a single spiracle. The sternum is the most
widely visible part of the abdomen, being a more or less sclerotized
segment. The abdomen itself does not have any appendages, but some
(for example, Mordellidae) have articulating sternal lobes.
Anatomy and physiology
A beetle's body systems
The digestive system of beetles is primarily adapted for a herbivorous
diet. Digestion takes place mostly in the anterior midgut, although in
predatory groups like the Carabidae, most digestion occurs in the crop
by means of midgut enzymes. In the Elateridae, the larvae are liquid
feeders that extraorally digest their food by secreting enzymes.
The alimentary canal basically consists of a short, narrow pharynx, a
widened expansion, the crop, and a poorly developed gizzard. This is
followed by the midgut, that varies in dimensions between species,
with a large amount of cecum, and the hindgut, with varying lengths.
There are typically four to six Malpighian tubules.
The nervous system in beetles contains all the types found in insects,
varying between different species, from three thoracic and seven or
eight abdominal ganglia which can be distinguished to that in which
all the thoracic and abdominal ganglia are fused to form a composite
Like most insects, beetles inhale air, for the oxygen it contains, and
exhale carbon dioxide, via a tracheal system. Air enters the body
through spiracles, and circulates within the haemocoel in a system of
tracheae and tracheoles, through whose walls the gases can diffuse.
Dytiscus spiracles (right) on upper side of abdomen, normally covered
by the elytra, are in contact with an air bubble when the beetle
Diving beetles, such as the Dytiscidae, carry a bubble of air with
them when they dive. Such a bubble may be contained under the elytra
or against the body by specialized hydrophobic hairs. The bubble
covers at least some of the spiracles, permitting air to enter the
tracheae. The function of the bubble is not only to contain a store
of air, but to act as a physical gill. The air that it traps is in
contact with oxygenated water, so as the animal's consumption depletes
the oxygen in the bubble, more oxygen can diffuse in to replenish
Carbon dioxide is more soluble in water than either oxygen or
nitrogen, so it readily diffuses out faster than in. Nitrogen is the
most plentiful gas in the bubble, and the least soluble, so it
constitutes a relatively static component of the bubble and acts as a
stable medium for respiratory gases to accumulate in and pass through.
Occasional visits to the surface are sufficient for the beetle to
re-establish the constitution of the bubble.
Like other insects, beetles have open circulatory systems, based on
hemolymph rather than blood. As in other insects, a segmented
tube-like heart is attached to the dorsal wall of the hemocoel. It has
paired inlets or ostia at intervals down its length, and circulates
the hemolymph from the main cavity of the haemocoel and out through
the anterior cavity in the head.
Different glands are specialized for different pheromones to attract
mates. Pheromones from species of
Rutelinae are produced from
epithelial cells lining the inner surface of the apical abdominal
segments; amino acid-based pheromones of
Melolonthinae are produced
from eversible glands on the abdominal apex. Other species produce
different types of pheromones. Dermestids produce esters, and species
Elateridae produce fatty acid-derived aldehydes and acetates. To
attract a mate, fireflies (Lampyridae) use modified fat body cells
with transparent surfaces backed with reflective uric acid crystals to
produce light by bioluminescence. Light production is highly
efficient, by oxidation of luciferin catalyzed by enzymes
(luciferases) in the presence of adenosine triphosphate (ATP) and
oxygen, producing oxyluciferin, carbon dioxide, and light.
Tympanal organs or hearing organs consist of a membrane (tympanum)
stretched across a frame backed by an air sac and associated sensory
neurons, are found in two families. Several species of the genus
Cicindela (Carabidae) have hearing organs on the dorsal surfaces of
their first abdominal segments beneath the wings; two tribes in the
Dynastinae (within the Scarabaeidae) have hearing organs just beneath
their pronotal shields or neck membranes. Both families are sensitive
to ultrasonic frequencies, with strong evidence indicating they
function to detect the presence of bats by their ultrasonic
Reproduction and development
Beetles are members of the superorder Endopterygota, and accordingly
most of them undergo complete metamorphosis. The typical form of
metamorphosis in beetles passes through four main stages: the egg, the
larva, the pupa, and the imago or adult. The larvae are commonly
called grubs and the pupa sometimes is called the chrysalis. In some
species, the pupa may be enclosed in a cocoon constructed by the larva
towards the end of its final instar. Some beetles, such as typical
members of the families
Meloidae and Rhipiphoridae, go further,
undergoing hypermetamorphosis in which the first instar takes the form
of a triungulin.
Punctate flower chafers (Neorrhina punctata, Scarabaeidae) mating
Some beetles have intricate mating behaviour.
is often important in locating a mate. Different species use different
pheromones. Scarab beetles such as the
Rutelinae use pheromones
derived from fatty acid synthesis, while other scarabs such as the
Melolonthinae use amino acids and terpenoids. Another way beetles find
mates is seen in the fireflies (Lampyridae) which are bioluminescent,
with abdominal light-producing organs. The males and females engage in
a complex dialogue before mating; each species has a unique
combination of flight patterns, duration, composition, and intensity
of the light produced.
Before mating, males and females may stridulate, or vibrate the
objects they are on. In the Meloidae, the male climbs onto the dorsum
of the female and strokes his antennae on her head, palps, and
antennae. In Eupompha, the male draws his antennae along his
longitudinal vertex. They may not mate at all if they do not perform
the precopulatory ritual. This mating behaviour may be different
amongst dispersed populations of the same species. For example, the
mating of a Russian population of tansy beetle (Chysolina graminis) is
preceded by an elaborate ritual involving the male tapping the
female's eyes, pronotum and antennae with its antennae, which is not
evident in the population of this species in the United Kingdom.
Competition can play a part in the mating rituals of species such as
burying beetles (Nicrophorus), the insects fighting to determine which
can mate. Many male beetles are territorial and fiercely defend their
territories from intruding males. In such species, the male often has
horns on the head or thorax, making its body length greater than that
of a female. Copulation is generally quick, but in some cases lasts
for several hours. During copulation, sperm cells are transferred to
the female to fertilize the egg.
The life cycle of the stag beetle includes three instars.
Essentially all beetles lay eggs, though some myrmecophilous
Aleocharinae and some
Chrysomelinae which live in mountains or the
subarctic are ovoviviparous, laying eggs which hatch almost
Beetle eggs generally have smooth surfaces and are soft,
Cupedidae have hard eggs. Eggs vary widely between species:
the eggs tend to be small in species with many instars (larval
stages), and in those that lay large numbers of eggs. A female may lay
from several dozen to several thousand eggs during her lifetime,
depending on the extent of parental care. This ranges from the simple
laying of eggs under a leaf, to the parental care provided by scarab
beetles, which house, feed and protect their young. The Attelabidae
roll leaves and lay their eggs inside the roll for protection.
The larva is usually the principal feeding stage of the beetle life
cycle. Larvae tend to feed voraciously once they emerge from their
eggs. Some feed externally on plants, such as those of certain leaf
beetles, while others feed within their food sources. Examples of
internal feeders are most
Buprestidae and longhorn beetles. The larvae
of many beetle families are predatory like the adults (ground beetles,
ladybirds, rove beetles). The larval period varies between species,
but can be as long as several years. The larvae of skin beetles
undergo a degree of reversed development when starved, and later grow
back to the previously attained level of maturity. The cycle can be
repeated many times (see Biological immortality). Larval
morphology is highly varied amongst species, with well-developed and
sclerotized heads, distinguishable thoracic and abdominal segments
(usually the tenth, though sometimes the eighth or ninth).
Scarabaeiform larva of Hercules beetle
Beetle larvae can be differentiated from other insect larvae by their
hardened, often darkened heads, the presence of chewing mouthparts,
and spiracles along the sides of their bodies. Like adult beetles, the
larvae are varied in appearance, particularly between beetle families.
Beetles with somewhat flattened, highly mobile larvae include the
ground beetles and rove beetles; their larvae are described as
campodeiform. Some beetle larvae resemble hardened worms with dark
head capsules and minute legs. These are elateriform larvae, and are
found in the click beetle (Elateridae) and darkling beetle
(Tenebrionidae) families. Some elateriform larvae of click beetles are
known as wireworms. Beetles in the
Scarabaeoidea have short, thick
larvae described as scarabaeiform, more commonly known as grubs.
All beetle larvae go through several instars, which are the
developmental stages between each moult. In many species, the larvae
simply increase in size with each successive instar as more food is
consumed. In some cases, however, more dramatic changes occur. Among
certain beetle families or genera, particularly those that exhibit
parasitic lifestyles, the first instar (the planidium) is highly
mobile to search out a host, while the following instars are more
sedentary and remain on or within their host. This is known as
hypermetamorphosis; it occurs in the Meloidae, Micromalthidae, and
Ripiphoridae. The blister beetle
Epicauta vittata (Meloidae), for
example, has three distinct larval stages. Its first stage, the
triungulin, has longer legs to go in search of the eggs of
grasshoppers. After feeding for a week it moults to the second stage,
called the caraboid stage, which resembles the larva of a carabid
beetle. In another week it moults and assumes the appearance of a
scarabaeid larva – the scarabaeidoid stage. Its penultimate larval
stage is the pseudo-pupa or the coarcate larva, which will overwinter
and pupate until the next spring.
The larval period can vary widely. A fungus feeding staphylinid
Phanerota fasciata undergoes three moults in 3.2 days at room
temperature while Anisotoma sp. (Leiodidae) completes its larval stage
in the fruiting body of slime mold in 2 days and possibly represents
the fastest growing beetles. Dermestid beetles, Trogoderma inclusum
can remain in an extended larval state under unfavourable conditions,
even reducing their size between moults. A larva is reported to have
survived for 3.5 years in an enclosed container.
Pupa and adult
The ivory-marked beetle, Eburia quadrigeminata, may live up to 40
years inside the hardwoods on which the larva feeds.
As with all endopterygotes, beetle larvae pupate, and from these pupae
emerge fully formed, sexually mature adult beetles, or imagos. Pupae
never have mandibles (they are adecticous). In most pupae, the
appendages are not attached to the body and are said to be exarate; in
a few beetles (Staphylinidae,
Ptiliidae etc.) the appendages are fused
with the body (termed as obtect pupae).
Adults have extremely variable lifespans, from weeks to years,
depending on the species. Some wood-boring beetles can have
extremely long life-cycles. It is believed that when furniture or
house timbers are infested by beetle larvae, the timber already
contained the larvae when it was first sawn up. A birch bookcase 40
years old released adult
Eburia quadrigeminata (Cerambycidae), while
Buprestis aurulenta and other
Buprestidae have been documented as
emerging as much as 51 years after manufacture of wooden items.
Photinus pyralis, firefly, in flight
The elytra allow beetles to both fly and move through confined spaces,
doing so by folding the delicate wings under the elytra while not
flying, and folding their wings out just before take off. The
unfolding and folding of the wings is operated by muscles attached to
the wing base; as long as the tension on the radial and cubital veins
remains, the wings remain straight. In some day-flying species (for
example, Buprestidae, Scarabaeidae), flight does not include large
amounts of lifting of the elytra, having the metathorac wings extended
under the lateral elytra margins. The altitude reached by beetles
in flight varies. One study investigating the flight altitude of the
Coccinella septempunctata and
Harmonia axyridis using
radar showed that, whilst the majority in flight over a single
location were at 150–195 m above ground level, some reached
altitudes of over 1100 m.
Many rove beetles have greatly reduced elytra, and while they are
capable of flight, they most often move on the ground: their soft
bodies and strong abdominal muscles make them flexible, easily able to
wriggle into small cracks.
Aquatic beetles use several techniques for retaining air beneath the
water's surface. Diving beetles (Dytiscidae) hold air between the
abdomen and the elytra when diving.
Hydrophilidae have hairs on their
under surface that retain a layer of air against their bodies. Adult
crawling water beetles use both their elytra and their hind coxae (the
basal segment of the back legs) in air retention, while whirligig
beetles simply carry an air bubble down with them whenever they
Beetles have a variety of ways to communicate, including the use of
pheromones. The mountain pine beetle emits a pheromone to attract
other beetles to a tree. The mass of beetles are able to overcome the
chemical defenses of the tree. After the tree's defenses have been
exhausted, the beetles emit an anti-aggregation pheromone. The species
can stridulate to communicate.
A dung beetle rolling dung
Parental care is found in a few species of beetle, perhaps for
protection against adverse conditions and predators. The rove
Bledius spectabilis lives in salt marshes, so the eggs and
larvae are endangered by the rising tide. The maternal beetle patrols
the eggs and larvae, burrowing to keep them from flooding and
asphyxiating, and protects them from the predatory carabid beetle
Dicheirotrichus gustavi and from the parasitoidal wasp Barycnemis
blediator, which kills some 15% of the larvae.
Burying beetles are attentive parents, and participate in cooperative
care and feeding of their offspring. Both parents work to bury small
animal carcass to serve as a food resource for their young and build a
brood chamber around it. The parents prepare the carcass and protect
it from competitors and from early decomposition. After their eggs
hatch, the parents keep the larvae clean of fungus and bacteria and
help the larvae feed by regurgitating food for them.
Some dung beetles provide parental care, collecting herbivore dung and
laying eggs within that food supply, an instance of mass provisioning.
Some species do not leave after this stage, but remain to safeguard
Most species of beetles do not display parental care behaviors after
the eggs have been laid.
Eusociality involves cooperative brood care (including brood care of
offspring from other individuals), overlapping generations within a
colony of adults, and a division of labour into reproductive and
non-reproductive groups. Few organisms outside
this behavior; the only beetle to do so is the weevil Austroplatypus
incompertus. This Australian species lives in horizontal networks
of tunnels, in the heartwood of
Eucalyptus trees. It is one of more
than 300 species of wood-boring Ambrosia beetles which distribute the
spores of ambrosia fungi. The fungi grow in the beetles' tunnels,
providing food for the beetles and their larvae; female offspring
remain in the tunnels and maintain the fungal growth, probably never
reproducing. Cooperative brood care is also found in the bess
beetles (Passalidae) where the larvae feed on the semi-digested faeces
of the adults.
Hycleus sp. (Meloidae) feeding on the petals of Ipomoea carnea
Beetles are able to exploit a wide diversity of food sources available
in their many habitats. Some are omnivores, eating both plants and
animals. Other beetles are highly specialized in their diet. Many
species of leaf beetles, longhorn beetles, and weevils are very
host-specific, feeding on only a single species of plant. Ground
beetles and rove beetles (Staphylinidae), among others, are primarily
carnivorous and catch and consume many other arthropods and small
prey, such as earthworms and snails. While most predatory beetles are
generalists, a few species have more specific prey requirements or
Decaying organic matter is a primary diet for many species. This can
range from dung, which is consumed by coprophagous species (such as
certain scarab beetles in the Scarabaeidae), to dead animals, which
are eaten by necrophagous species (such as the carrion beetles,
Silphidae). Some beetles found in dung and carrion are in fact
predatory. These include members of the
Histeridae and Silphidae,
preying on the larvae of coprophagous and necrophagous insects.
Many beetles feed under bark, some feed on wood while others feed on
fungi growing on wood or leaf-litter. Some beetles have special
mycangia, structures for the transport of fungal spores.
A camouflaged longhorn beetle, Ecyrus dasycerus
Beetles, both adults and larvae, are the prey of many animal predators
including mammals from bats to rodents, birds, lizards, amphibians,
fishes, dragonflies, robberflies, reduviid bugs, ants, other beetles,
and spiders. Beetles use a variety of anti-predator
adaptations to defend themselves. These include camouflage and mimicry
against predators that hunt by sight, toxicity, and defensive
Further information: Camouflage
Camouflage is common and widespread among beetle families, especially
those that feed on wood or vegetation, such as leaf beetles
(Chrysomelidae, which are often green) and weevils. In some species,
sculpturing or various coloured scales or hairs cause beetles such as
the avocado weevil Heilipus apiatus to resemble bird dung or other
inedible objects. Many beetles that live in sandy environments
blend in with the coloration of that substrate.
Mimicry and aposematism
Clytus arietis (Cerambycidae), a Batesian mimic of wasps
Mimicry and Aposematism
Some longhorn beetles (Cerambycidae) are effective Batesian mimics of
wasps. Beetles may combine coloration with behavioural mimicry, acting
like the wasps they already closely resemble. Many other beetles,
including ladybirds, blister beetles, and lycid beetles secrete
distasteful or toxic substances to make them unpalatable or poisonous,
and are often aposematic, where bright or contrasting coloration warn
off predators; many beetles and other insects mimic these chemically
Blister beetles such as
Hycleus have brilliant aposematic coloration,
warning of their toxicity.
The bloody-nosed beetle, Timarcha tenebricosa, defending itself by
releasing a droplet of noxious red liquid (base of leg, on right)
Chemical defense is important in some species, usually being
advertised by bright aposematic colours. Some
Tenebrionidae use their
posture for releasing noxious chemicals to warn off predators.
Chemical defences may serve purposes other than just protection from
vertebrates, such as protection from a wide range of microbes. Some
species sequester chemicals from the plants they feed on,
incorporating them into their own defenses.
Other species have special glands to produce deterrent chemicals. The
defensive glands of carabid ground beetles produce a variety of
hydrocarbons, aldehydes, phenols, quinones, esters, and acids released
from an opening at the end of the abdomen. African carabid beetles
Thermophilum is sometimes
included within Anthia) employ the same chemicals as ants: formic
acid. Bombardier beetles have well-developed pygidial glands that
empty from the sides of the intersegment membranes between the seventh
and eighth abdominal segments. The gland is made of two containing
chambers, one for hydroquinones and hydrogen peroxide, the other
holding hydrogen peroxide and catalase enzymes. These chemicals mix
and result in an explosive ejection, reaching a temperature of around
100 °C (212 °F), with the breakdown of hydroquinone to
hydrogen, oxygen, and quinone. The oxygen propels the noxious chemical
spray as a jet that can be aimed accurately at predators.
Large ground-dwelling beetles such as Carabidae, the rhinoceros beetle
and the longhorn beetles defend themselves using strong mandibles, or
heavily sclerotised (armored) spines or horns to deter or fight off
predators. Many species of weevil that feed out in the open on
leaves of plants react to attack by employing a drop-off reflex. Some
combine it with thanatosis, in which they close up their appendages
and "play dead". The click beetles (Elateridae) can suddenly
catapult themselves out of danger by releasing the energy stored by a
click mechanism, which consists of a stout spine on the prosternum and
a matching groove in the mesosternum.
A few species of beetles are ectoparasitic on mammals. One such
species, Platypsyllus castoris, parasitises beavers (Castor spp.).
This beetle lives as a parasite both as a larva and as an adult,
feeding on epidermal tissue and possibly on skin secretions and wound
exudates. They are strikingly flattened dorsoventrally, no doubt as an
adaptation for slipping between the beavers' hairs. They are wingless
and eyeless, as are many other ectoparasites. Others are
kleptoparasites of other invertebrates, such as the small hive beetle
(Aethina tumida) that infests honey bee nests, while many species
are parasitic inquilines or commensal in the nests of ants. A few
groups of beetles are primary parasitoids of other insects, feeding
off of, and eventually killing their hosts.
Rose chafer pollinating a labiate flower
Beetle-pollinated flowers are usually large, greenish or off-white in
color, and heavily scented. Scents may be spicy, fruity, or similar to
decaying organic material. Beetles were most likely the first insects
to pollinate flowers. Most beetle-pollinated flowers are flattened or
dish-shaped, with pollen easily accessible, although they may include
traps to keep the beetle longer. The plants' ovaries are usually well
protected from the biting mouthparts of their pollinators. The beetle
families that habitually pollinate flowers are the Buprestidae,
Cantharidae, Carambycidae, Cleridae, Dermestidae, Lycidae, Melyridae,
Mordellidae, Nitidulidae and Scarabeidae. Beetles may be
particularly important in some parts of the world such as semiarid
areas of southern Africa and southern California and the montane
KwaZulu-Natal in South Africa.
1: Adult ambrosia beetle burrows into wood and lays eggs, carrying
fungal spores in its mycangia.
Larva feeds on fungus, which digests wood, removing toxins, to
Mutualism is well known in a few beetles, such as the ambrosia beetle,
which partners with fungi to digest the wood of dead trees. The
beetles excavate tunnels in dead trees in which they cultivate fungal
gardens, their sole source of nutrition. After landing on a suitable
tree, an ambrosia beetle excavates a tunnel in which it releases
spores of its fungal symbiont. The fungus penetrates the plant's xylem
tissue, digests it, and concentrates the nutrients on and near the
surface of the beetle gallery, so the weevils and the fungus both
benefit. The beetles cannot eat the wood due to toxins, and uses its
relationship with fungi to help overcome the defenses of its host tree
in order to provide nutrition for their larvae. Chemically
mediated by a bacterially produced polyunsaturated peroxide, this
mutualistic relationship between the beetle and the fungus is
Tolerance of extreme environments
The fogstand beetle of the Namib Desert,
Stenocara gracilipes is able
to survive by collecting water from fog on its back.
Insect thermoregulation and
Insect winter ecology
About 90% of beetle species enter a period of adult diapause, a quiet
phase with reduced metabolism to tide unfavourable environmental
conditions. Adult diapause is the most common form of diapause in
Coleoptera. To endure the period without food (often lasting many
months) adults prepare by accumulating reserves of lipids, glycogen,
proteins and other substances needed for resistance to future
hazardous changes of environmental conditions. This diapause is
induced by signals heralding the arrival of the unfavourable season;
usually the cue is photoperiodic. Short (decreasing) day length serves
as a signal of approaching winter and induces winter diapause
(hibernation). A study of hibernation in the Arctic beetle
Pterostichus brevicorni showed that the body fat levels of adults were
highest in autumn with the alimentary canal filled with food, but
empty by the end of January. This loss of body fat was a gradual
process, occurring in combination with dehydration.
All insects are poikilothermic, so the ability of a few beetles
to live in extreme environments depends on their resilience to
unusually high or low temperatures. The bark beetle Pityogenes
chalcographus can survive −39°C whilst overwintering beneath tree
bark; the Alaskan beetle
Cucujus clavipes puniceus is able to
withstand −58°C; its larvae may survive −100°C. At these
low temperatures, the formation of ice crystals in internal fluids is
the biggest threat to survival to beetles, but this is prevented
through the production of antifreeze proteins that stop water
molecules from grouping together. The low temperatures experienced by
Cucujus clavipes can be survived through their deliberate dehydration
in conjunction with the antifreeze proteins. This concentrates the
antifreezes several fold. The hemolymph of the mealworm beetle
Tenebrio molitor contains several antifreeze proteins. The
Upis ceramboides can survive −60 °C: its
cryoprotectants are xylomannan, a molecule consisting of a sugar bound
to a fatty acid, and the sugar-alcohol, threitol.
Conversely, desert dwelling beetles are adapted to tolerate high
temperatures. For example, the Tenebrionid beetle Onymacris
rugatipennis can withstand 50°C. Tiger beetles in hot, sandy
areas are often whitish (for example, Habroscelimorpha dorsalis), to
reflect more heat than a darker colour would. These beetles also
exhibits behavioural adaptions to tolerate the heat: they are able to
stand erect on their tarsi to hold their bodies away from the hot
ground, seek shade, and turn to face the sun so that only the front
parts of their heads are directly exposed.
The fogstand beetle of the Namib Desert, Stenocara gracilipes, is able
to collect water from fog, as its elytra have a textured surface
combining hydrophilic (water-loving) bumps and waxy, hydrophobic
troughs. The beetle faces the early morning breeze, holding up its
abdomen; droplets condense on the elytra and run along ridges towards
their mouthparts. Similar adaptations are found in several other Namib
desert beetles such as Onymacris unguicularis.
Some terrestrial beetles that exploit shoreline and floodplain
habitats have physiological adaptations for surviving floods. In the
event of flooding, adult beetles may be mobile enough to move away
from flooding, but larvae and pupa often cannot. Adults of Cicindela
togata are unable to survive immersion in water, but larvae are able
to survive a prolonged period, up to 6 days, of anoxia during floods.
Anoxia tolerance in the larvae may have been sustained by switching to
anaerobic metabolic pathways or by reducing metabolic rate.
Anoxia tolerance in the adult Carabid beetle Pelophilia borealis was
tested in laboratory conditions and it was found that they could
survive a continuous period of up to 127 days in an atmosphere of
99.9% nitrogen at 0 °C.
Many beetle species undertake annual mass movements which are termed
as migrations. These include the pollen beetle Meligethes aeneus
and many species of coccinellids. These mass movements may also
be opportunistic, in search of food, rather than seasonal. A 2008
study of an unusually large outbreak of Mountain Pine Beetle
(Dendroctonus ponderosae) in
British Columbia found that beetles were
capable of flying 30–110 km per day in densities of up to 18,
600 beetles per hectare.
Relationship to humans
In ancient cultures
A scarab in the Valley of the Kings
Main article: Scarab (artifact)
Several species of dung beetle, especially the sacred scarab,
Scarabaeus sacer, were revered in Ancient Egypt. The
hieroglyphic image of the beetle may have had existential, fictional,
or ontologic significance. Images of the scarab in bone, ivory,
stone, Egyptian faience, and precious metals are known from the Sixth
Dynasty and up to the period of Roman rule. The scarab was of prime
significance in the funerary cult of ancient Egypt. The scarab
was linked to Khepri, the god of the rising sun, from the supposed
resemblance of the rolling of the dung ball by the beetle to the
rolling of the sun by the god. Some of ancient Egypt's neighbors
adopted the scarab motif for seals of varying types. The best-known of
these are the Judean LMLK seals, where eight of 21 designs contained
scarab beetles, which were used exclusively to stamp impressions on
storage jars during the reign of Hezekiah. Beetles are mentioned
as a symbol of the sun, as in ancient Egypt, in Plutarch's 1st century
Greek Magical Papyri of the 2nd century BC to the
5th century AD describe scarabs as an ingredient in a spell.
Pliny the Elder
Pliny the Elder discusses beetles in his Natural History,
describing the stag beetle: "Some insects, for the preservation of
their wings, are covered with a erust (elytra) – the beetle, for
instance, the wing of which is peculiarly fine and frail. To these
insects a sting has been denied by Nature; but in one large kind we
find horns of a remarkable length, two-pronged at the extremities, and
forming pincers, which the animal closes when it is its intention to
bite." The stag beetle is recorded in a Greek myth by
Nicander and recalled by
Antoninus Liberalis in which Cerambus[b] is
turned into a beetle: "He can be seen on trunks and has hook-teeth,
ever moving his jaws together. He is black, long and has hard wings
like a great dung beetle". The story concludes with the comment
that the beetles were used as toys by young boys, and that the head
was removed and worn as a pendant.
Cotton boll weevil
About 75% of beetle species are phytophagous in both the larval and
adult stages. Many feed on economically important plants and stored
plant products, including trees, cereals, tobacco, and dried
fruits. Some, such as the boll weevil, which feeds on cotton buds
and flowers, can cause extremely serious damage to agriculture. The
boll weevil crossed the
Rio Grande near Brownsville, Texas, to enter
the United States from
Mexico around 1892, and had reached
Alabama by 1915. By the mid-1920s, it had entered all
cotton-growing regions in the US, traveling 40 to 160 miles
(60–260 km) per year. It remains the most destructive cotton
pest in North America.
Mississippi State University
Mississippi State University has estimated,
since the boll weevil entered the United States, it has cost cotton
producers about $13 billion, and in recent times about $300 million
The bark beetle, elm leaf beetle and the Asian longhorned beetle
(Anoplophora glabripennis) are among the species that attack elm
trees. Bark beetles (Scolytidae) carry
Dutch elm disease
Dutch elm disease as they move
from infected breeding sites to healthy trees. The disease has
devastated elm trees across Europe and North America.
Larvae of the Colorado potato beetle, Leptinotarsa decemlineata, a
serious crop pest
Some species of beetle have evolved immunity to insecticides. For
example, the Colorado potato beetle, Leptinotarsa decemlineata, is a
destructive pest of potato plants. Its hosts include other members of
the Solanaceae, such as nightshade, tomato, eggplant and capsicum, as
well as the potato. Different populations have between them developed
resistance to all major classes of insecticide. The Colorado
potato beetle was evaluated as a tool of entomological warfare during
World War II, the idea being to use the beetle and its larvae to
damage the crops of enemy nations. Germany tested its Colorado
potato beetle weaponisation program south of Frankfurt, releasing
The death watch beetle, Xestobium rufovillosum (Anobiidae), is a
serious pest of older wooden buildings in Europe. It attacks hardwoods
such as oak and chestnut, always where some fungal decay has taken or
is taking place. The actual introduction of the pest into buildings is
thought to take place at the time of construction.
Other pests include the coconut hispine beetle, Brontispa longissima,
which feeds on young leaves, seedlings and mature coconut trees,
causing serious economic damage in the Philippines. The mountain
pine beetle is a destructive pest of mature or weakened lodgepole
pine, sometimes affecting large areas of Canada.
As beneficial resources
Coccinella septempunctata, a predatory beetle beneficial to
Beetles can be beneficial to human economics by controlling the
populations of pests. The larvae and adults of some species of lady
beetles (Coccinellidae) feed on aphids that are pests. Other lady
beetles feed on scale insects, whitefly and mealybugs. If normal
food sources are scarce, they may feed on small caterpillars, young
plant bugs, or honeydew and nectar. Ground beetles (Carabidae)
are common predators of many insect pests, including fly eggs,
caterpillars, and wireworms. Ground beetles can help to control
weeds by eating their seeds in the soil, reducing the need for
herbicides to protect crops. The effectiveness of some species in
reducing certain plant populations has resulted in the deliberate
introduction of beetles in order to control weeds. For example, the
Zygogramma is native to North America but has been used to
Parthenium hysterophorus in India and Ambrosia artemisiifolia
Dung beetles (Scarabidae) have been successfully used to reduce the
populations of pestilent flies, such as
Musca vetustissima and
Haematobia exigua which are serious pests of cattle in Australia.
The beetles make the dung unavailable to breeding pests by quickly
rolling and burying it in the soil, with the added effect of improving
soil fertility, tilth, and nutrient cycling. The Australian Dung
Beetle Project (1965–1985), introduced species of dung beetle to
Australia from South Africa and Europe to reduce populations of Musca
vetustissima, following successful trials of this technique in
American Institute of Biological Sciences reports
that dung beetles save the United States cattle industry an estimated
US$380 million annually through burying above-ground livestock
Dermestidae are often used in taxidermy and in the preparation of
scientific specimens, to clean soft tissue from bones. Larvae
feed on and remove cartilage along with other soft tissue.
Mealworms in a bowl for human consumption
Main article: Entomophagy
Beetles are the most widely eaten insects, with about 344 species used
as food, usually at the larval stage. The mealworm (the larva of
the darkling beetle) and the rhinoceros beetle are among the species
As biodiversity indicators
Due to their habitat specificity, many species of beetles have been
suggested as suitable as indicators, their presence, numbers, or
absence providing a measure of habitat quality. Predatory beetles such
as the tiger beetles (Cicindelidae) have found scientific use as an
indicator taxon for measuring regional patterns of biodiversity. They
are suitable for this as their taxonomy is stable; their life history
is well described; they are large and simple to observe when visiting
a site; they occur around the world in many habitats, with species
specialised to particular habitats; and their occurrence by species
accurately indicates other species, both vertebrate and
invertebrate. According to the habitats, many other groups such
as the rove beetles in human-modified habitats, dung beetles in
savannas and saproxylic beetles in forests have been
suggested as potential indicator species.
In art and adornment
Beetlewing and Live insect jewelry
Zopheridae in jewellery at the Texas A&M University Insect
Pendant watch in shape of beetle, Switzerland 1850-1900 gold, diamond,
Many beetles have beautiful and durable elytra that have been used as
material in arts, with beetlewing the best example. Sometimes,
they are incorporated into ritual objects for their religious
significance. Whole beetles, either as-is or encased in clear plastic,
are made into objects ranging from cheap souvenirs such as key chains
to expensive fine-art jewellery. In parts of Mexico, beetles of the
Zopherus are made into living brooches by attaching costume
jewelry and golden chains, which is made possible by the incredibly
hard elytra and sedentary habits of the genus.
Fighting beetles are used for entertainment and gambling. This sport
exploits the territorial behavior and mating competition of certain
species of large beetles. In the
Chiang Mai district of northern
Xylotrupes rhinoceros beetles are caught in the wild
and trained for fighting. Females are held inside a log to stimulate
the fighting males with their pheromones. These fights may be
competitive and involve gambling both money and property. In
South Korea the
Dytiscidae species Cybister tripunctatus is used in a
Beetles are sometimes used as instruments: the Onabasulu of Papua New
Guinea historically used the weevil
Rhynchophorus ferrugineus as a
musical instrument by letting the human mouth serve as a variable
resonance chamber for the wing vibrations of the live adult
Some species of beetle are kept as pets, for example diving beetles
(Dytiscidae) may be kept in a domestic fresh water tank.
"Remarkable Beetles Found at Simunjon, Borneo".[c] A few of the 2000
species of beetle collected by
Alfred Russel Wallace
Alfred Russel Wallace in Borneo
Japan the practice of keeping horned rhinoceros beetles
(Dynastinae) and stag beetles (Lucanidae) is particularly popular
amongst young boys. Such is the popularity in
Japan that vending
machines dispensing live beetles were developed in 1999, each holding
up to 100 stag beetles.
As things to collect
Beetle collecting became extremely popular in the Victorian era.
Alfred Russel Wallace
Alfred Russel Wallace collected (by his own count) a
total of 83,200 beetles during the eight years described in his 1869
book The Malay Archipelago, including 2,000 species new to
As inspiration for technologies
Further information: Biomimetics
Several coleopteran adaptations have attracted interest in biomimetics
with possible commercial applications. The bombardier beetle's
powerful repellent spray has inspired the development of a fine mist
spray technology, claimed to have a low carbon impact compared to
aerosol sprays. Moisture harvesting behavior by the Namib desert
beetle (Stenocara gracilipes) has inspired a self-filling water bottle
which utilises hydrophilic and hydrophobic materials to benefit people
living in dry regions with no regular rainfall.
Living beetles have been used as cyborgs. A Defense Advanced Research
Projects Agency funded project implanted electrodes into Mecynorhina
torquata beetles, allowing them to be remotely controlled via a radio
receiver held on its back, as proof-of-concept for surveillance
work. Similar technology has been applied to enable a human
operator to control the free-flight steering and walking gaits of
Mecynorhina torquata as well as graded turning and backward walking of
Since beetles form such a large part of the world's biodiversity,
their conservation is important, and equally, loss of habitat and
biodiversity is essentially certain to impact on beetles.
Unfortunately, with exceptions for groups such as ladybirds and
fireflies, beetles as a whole have a less positive image than other
insect groups like the butterflies: many are indeed pests, and others
arouse disgust. Many species of beetles have very specific habitats
and long life cycles that make them vulnerable. Some species are
highly threatened while others are already feared extinct. Island
species tend to be more susceptible as in the case of Helictopleurus
undatus of Madagascar which is thought to have gone extinct during the
late 20th century. Conservationists have attempted to arouse a
liking for beetles with flagship species like the stag beetle, Lucanus
cervus, and tiger beetles (Cicindelidae). In
Japan the Genji
firefly, Luciola cruciata, is extremely popular, and in South Africa
the Addo elephant dung beetle offers promise for broadening ecotourism
beyond the big five tourist mammal species. Popular dislike of pest
beetles, too, can be turned into public interest in insects, as can
unusual ecological adaptations of species like the fairy shrimp
hunting beetle, Cicinis bruchi.
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Wikispecies has information related to Coleoptera
The Wikibook Dichotomous Key has a page on the topic of: Coleoptera
Coleoptera from the Tree of Life Web Project
(in German) Käfer der Welt
Beetles – Coleoptera
Archaeognatha (jumping bristletails)
Thysanura (Zygentoma) (silverfish, firebrats)
Odonata (dragonflies, damselflies)
Phasmatodea (stick and leaf insects)
Notoptera (ice-crawlers, gladiators)
Orthoptera (crickets, wetas, grasshoppers, locusts)
Zoraptera (angel insects)
Blattodea (cockroaches, termites)
Psocodea (barklice, lice)
Hemiptera (cicadas, aphids, true bugs)
Hymenoptera (sawflies, wasps, ants, bees)
Strepsiptera (twisted-winged parasites)
Megaloptera (alderflies, dobsonflies, fishflies)
Neuroptera (net-winged insects: lacewings, mantidflies, antlions)
Mecoptera (scorpionflies) + Siphonaptera (fleas)
Diptera (gnats, mosquitoes, flies)
Lepidoptera (moths, butterflies)
Four most speciose orders are marked in bold
Italic are paraphyletic groups
Based on Sasaki et al. (2013)
Extinct incertae sedis families and genera are marked in italic
Extant Coleoptera families
Crowsoniellidae (Crowsoniella relicta)
Cupedidae (reticulated beetles)
Jurodidae (Sikhotealinia zhiltzovae)
Micromalthidae (telephone-pole beetle)
Amphizoidae (trout-stream beetles)
Carabidae (ground beetles)
Dytiscidae (predaceous diving beetles)
Gyrinidae (whirligig beetles)
Haliplidae (crawling water beetles)
Meruidae (Meru phyllisae)
Noteridae (burrowing water beetles)
Rhysodidae (wrinkled bark beetles)
Trachypachidae (false ground beetles)
Hydroscaphidae (skiff beetles)
Anobiidae (furniture beetles, death watch beetles, spider beetles)
Bostrichidae (auger beetles)
Dermestidae (skin beetles)
Jacobsoniidae (Jacobson's beetles)
Nosodendridae (wounded-tree beetles)
Derodontidae (tooth-necked fungus beetles)
Cerambycidae (longhorn beetles)
Chrysomelidae (leaf beetles)
Acanthocnemidae (Acanthocnemus nigricans)
Cleridae (checkered beetles)
Melyridae (soft-wing flower beetles)
Phloiophilidae (Phloiophilus edwardsi)
Trogossitidae (bark-gnawing beetles)
Biphyllidae (false skin beetles)
Bothrideridae (dry bark beetles)
Byturidae (fruitworm beetles)
Cerylonidae (minute bark beetles)
Coccinellidae (lady beetles, or God's cows)
Corylophidae (minute fungus beetles)
Cryptophagidae (silken fungus beetles)
Cucujidae (flat bark beetles)
Endomychidae (handsome fungus beetles)
Erotylidae (pleasing fungus beetles)
Kateretidae (short-winged flower beetles)
Laemophloeidae (lined flat bark beetles)
Lamingtoniidae (Lamingtonium binnaberrense)
Latridiidae (minute brown scavenger beetles)
Monotomidae (root-eating beetles)
Nitidulidae (sap beetles)
Passandridae (parasitic flat bark beetles)
Phalacridae (shining flower beetles)
Silvanidae (silvanid flat bark beetles)
Smicripidae (palmetto beetles)
Sphindidae (dry-fungus beetles)
Anthribidae (fungus weevils)
Attelabidae (leaf-rolling weevils)
Belidae (primitive weevils)
Brentidae (straight snout weevils, New York weevil)
Curculionidae (true weevils, bark beetles, ambrosia beetles)
Nemonychidae (pine flower weevils)
Lymexylidae (ship-timber beetles)
Aderidae (ant-like leaf beetles)
Anthicidae (ant-like flower beetles)
Archeocrypticidae (cryptic fungus beetles)
Boridae (conifer bark beetles)
Ciidae (minute tree-fungus beetles)
Melandryidae (false darkling beetles)
Meloidae (blister beetles)
Mordellidae (tumbling flower beetles)
Mycetophagidae (hairy fungus beetles)
Mycteridae (palm and flower beetles)
Oedemeridae (false blister beetle)
Perimylopidae, or Promecheilidae
Prostomidae (jugular-horned beetles)
Pyrochroidae (fire-coloured beetles)
Pythidae (dead log bark beetles)
Ripiphoridae (wedge-shaped beetles)
Salpingidae (narrow-waisted bark beetles)
Scraptiidae (false flower beetles)
Stenotrachelidae (false longhorn beetles)
Synchroidae (synchroa bark beetles)
Tenebrionidae (darkling beetles)
Tetratomidae (polypore fungus beetles)
Zopheridae (ironclad beetles, cylindrical bark beetles)
Buprestidae (jewel beetles, or metallic wood-boring beetles)
Byrrhidae (pill beetles)
Callirhipidae (cedar beetles)
Chelonariidae (turtle beetles)
Dryopidae (long-toed water beetles)
Elmidae (riffle beetles)
Eulichadidae (forest stream beetles)
Heteroceridae (variegated mud-loving beetles)
Limnichidae (minute mud beetles)
Lutrochidae (travertine beetles)
Psephenidae (water-penny beetles)
Dascillidae (soft bodied plant beetles)
Rhipiceridae (cicada beetle, cicada parasite beetles)
Artematopodidae (soft-bodied plant beetles)
Brachypsectridae (Texas beetles)
Cantharidae (soldier beetles)
Cerophytidae (rare click beetles)
Elateridae (click beetles)
Eucnemidae (false click beetles)
Lycidae (net-winged beetles)
Omethidae (false fireflies)
Phengodidae (glowworm beetles, long-lipped beetles)
Plastoceridae (Plastocerus angulosus)
Rhinorhipidae (Rhinorhipus tamborinensis)
Throscidae (false metallic wood-boring beetles)
Decliniidae (Declinia relicta)
Eucinetidae (plate-thigh beetles)
Diphyllostomatidae (false stag beetles)
Geotrupidae (dor beetles)
Glaphyridae (bumble bee scarab beetles)
Glaresidae (enigmatic scarab beetles)
Hybosoridae (scavenger scarab beetles)
Lucanidae (stag beetles)
Ochodaeidae (sand-loving scarab beetles)
Passalidae (betsy beetles)
Pleocomidae (rain beetles)
Trogidae (hide beetles)
Histeridae (clown beetles)
Sphaeritidae (false clown beetles)
Georissidae (minute mud-loving beetles)
Hydrophilidae (water scavenger beetles)
Agyrtidae (primitive carrion beetles)
Leiodidae (round fungus beetles)
Ptiliidae (feather-winged beetles)
Scydmaenidae (ant-like stone beetles)
Silphidae (carrion beetles)
Staphylinidae (rove beetles)
List of subgroups of the order Coleoptera
Insects in culture
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List of insect-inspired songs
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Colorado potato beetle
Cottony cushion scale
Western corn rootworm
Insect bites and stings
Insect sting allergy
House longhorn beetle
Home-stored product entomology
Alfred Russel Wallace
Hans Zinsser (Rats, Lice and History)
Lafcadio Hearn (
Living things in culture
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Crystal Palace Dinosaurs
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Rod of Asclepius
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In food processing
List of microbes
Microbes and Man
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Template:Insects in culture
Fauna Europaea: 10993