See subgroups of the order Coleoptera
BEETLES are a group of insects that form the order COLEOPTERA, in the
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 damage crops.
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
effective camouflage .
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
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
* 1 Etymology
* 2 Distribution and diversity
* 3 Evolution
* 4 Phylogeny
* 5 External morphology
* 5.1 Head
* 5.2 Thorax
* 5.3 Legs
* 5.4 Wings
Anatomy and physiology
* 6.3 Respiratory system
* 6.4 Circulatory system
* 6.5 Specialized organs
* 7 Reproduction and development
* 7.1 Mating
* 7.2 Life cycle
* 7.2.1 Egg
Pupa and adult
* 8 Behaviour
* 8.1 Locomotion
* 8.2 Communication
* 8.3 Parental care
* 8.5 Feeding
* 9 Ecology
* 9.1 Anti-predator adaptations
Mimicry and aposematism
* 9.1.3 Other defences
* 9.3 Pollination
* 9.4 Mutualism
* 9.5 Tolerance of extreme environments
* 9.6 Migration
* 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
* 11 Notes
* 12 References
* 12.1 Bibliography
* 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),
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 beetle Scydosella
musawasensis which may measure as little as 0.325 mm 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 (
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
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
Euramerica . The first discoveries from North America made in the
Wellington formation of Oklahoma were published in 2005 and 2008.
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 . Further
records are known from 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
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 mutualistic relationship.
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
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 basin.
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 fly
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 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 suborders.
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 beetle's legs).
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 hind leg.
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
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
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 ago (mya).
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 mandibulate mouthparts with the mandibles mobile in a
horizontal plane. The mouthparts are rarely suctorial and are
sometimes reduced and the maxillae always bear palps. The antennae
usually have 11 or fewer segments except in some groups like the
Rhipiceridae and Cerambycidae. 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 extant beetles.
Unlike the adults, beetle larvae can often be confused with larvae 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 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
Dermestidae ), some rove beetles (
Omaliinae ), and the
Polyphylla fullo has distinctive fan-like
antennae , one of several distinct forms for the appendages among
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 abdomen.
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
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
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
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
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
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
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
Dytiscus spiracles (right) on upper side of abdomen,
normally covered by the elytra, are in contact with an air bubble when
the beetle dives.
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 it.
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
Rhipiphoridae , go further,
undergoing hypermetamorphosis in which the first instar takes the form
of a triungulin .
Punctate flower chafers (Neorrhina punctata, Scarabaeidae)
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
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
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
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 dive.
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 beetle
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 their offspring.
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
Passalidae ) where the larvae feed on the semi-digested
faeces of the adults.
Hycleus sp. (
Meloidae ) feeding on the petals of
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
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 ,
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 behaviour.
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
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 protected species. 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 (for example,
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
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
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 grasslands
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 coevolved .
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. Further
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
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 Alaskan beetle
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
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
RELATIONSHIP TO HUMANS
IN ANCIENT CULTURES
A scarab in the
Valley of the Kings
Valley of the Kings Main article: Scarab
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
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
Hezekiah . Beetles are mentioned as a symbol of the sun, as
in ancient Egypt, in
Plutarch 's 1st century
Moralia . The 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
Antoninus Liberalis in which
Cerambus 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
Cotton boll weevil
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 southeastern
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 per year.
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
Colorado potato beetle
Colorado potato beetle , Leptinotarsa decemlineata, a serious crop
Some species of beetle have evolved immunity to insecticides. For
Colorado potato beetle
Colorado potato beetle , Leptinotarsa decemlineata, is a
destructive pest of potato plants. Its hosts include other members of
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
Colorado potato beetle was evaluated as a tool of entomological
World War II
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
Colorado potato beetle weaponisation program south of Frankfurt ,
releasing 54,000 beetles.
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 pest 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
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 genus Zygogramma
is native to North America but has been used to control Parthenium
hysterophorus in India and
Ambrosia artemisiifolia in Russia.
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
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
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 Hawaii
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 feces.
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:
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
IN ART AND ADORNMENT
Live insect jewelry
jewellery at the Texas A"> "Remarkable Beetles Found at Simunjon,
Borneo". A few of the 2000 species of beetle collected by Alfred
Russel Wallace in
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
The Malay Archipelago , including 2,000 species new to science.
AS INSPIRATION FOR TECHNOLOGIES
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.
Technological adaptations have also utilised living beetles as
cyborgs . For example, a Defense Advanced Research Projects Agency
funded project implanted electrodes into adult male Rhinoceros
beetles, allowing them to be remotely controlled via a radio receiver
held on its back. Electrical signals delivered via the electrodes
command the insect to take off, turn left or right, or hover in
midflight. The research was intended to suggest proof-of-concept for
surveillance purposes. Similar technology has been applied to the
Mecynorhina torquata to stimulate different walking
gaits and lengths of a live beetle controllable by a human operator.
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
* Coleoptera Atlas
* Beetles – Coleoptera
* Wd : Q22671
* ADW : Coleoptera
BugGuide : 60
* EoL : 345
* GBIF : 1470
* ITIS : 109216