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By clicking any link on this page you are giving your consent to our Privacy Policy and Cookies Policy. Aquatic beetles including the Dytiscidae diving 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. 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.

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.

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.

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.

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.

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.

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 structure.

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.

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 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.

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 of Elateridae produce fatty acid-derived aldehydes and acetates.

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.

Both families are sensitive to ultrasonic frequencies, with strong evidence indicating they function to detect the presence of bats by their ultrasonic echolocation.

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.

Some beetles have intricate mating behaviour. Pheromone communication 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.

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.

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 immediately.

Beetle eggs generally have smooth surfaces and are soft, though the 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. 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. 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. 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. 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.

Adults have extremely variable lifespans, from weeks to years, depending on the species. 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.

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 takeoff.

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.

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.

This species can stridulate to communicate, [92] but others may use sound to defend themselves when attacked. Parental care is found in a few families [94] of beetle, perhaps for protection against adverse conditions and predators.

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.

Subsociality, where females guard their offspring, is well-documented in two families of Chrysomelidae, Cassidinae and Chrysomelinae.

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.

It is one of more than species of wood-boring Ambrosia beetles which distribute the spores of ambrosia fungi. 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.

Some larger beetles or their larvae may consume vertebrates, particularly amphibians and fish. 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.

Some beetles have special mycangia , structures for the transport of fungal spores. 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.

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.

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.

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, Anthia employ the same chemicals as ants: formic acid. The gland is made of two containing chambers, one for hydroquinones and hydrogen peroxide , the other holding hydrogen peroxide and catalase enzymes.

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.

Some combine it with thanatosis , in which they close up their appendages and "play dead". 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.

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. 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 , Cerambycidae , Cleridae , Dermestidae , Lycidae , Melyridae , Mordellidae , Nitidulidae and Scarabaeidae.

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.

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. 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 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. Conversely, desert dwelling beetles are adapted to tolerate high temperatures. 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.

Many beetle species undertake annual mass movements which are termed as migrations. These include the pollen beetle Meligethes aeneus [] and many species of coccinellids.

Several species of dung beetle, especially the sacred scarab, Scarabaeus sacer , were revered in Ancient Egypt. The scarab was of prime significance in the funerary cult of ancient Egypt.

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.

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.

He is black, long and has hard wings like a great dung beetle". Many feed on economically important plants and stored plant products, including trees, cereals, tobacco, and dried fruits.

The boll weevil crossed the Rio Grande near Brownsville , Texas , to enter the United States from Mexico around , [] and had reached southeastern Alabama by It remains the most destructive cotton pest in North America.

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 as they move from infected breeding sites to healthy trees. The disease has devastated elm trees across Europe and North America.

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 death watch beetle , Xestobium rufovillosum Ptinidae , 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.

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. 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 Australia.

The Dermestidae are often used in taxidermy and in the preparation of scientific specimens, to clean soft tissue from bones.

Beetles are the most widely eaten insects, with about species used as food, usually at the larval stage. 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.

Many beetles have beautiful and durable elytra that have been used as material in arts, with beetlewing the best example.

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 genus 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 Thailand, male 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.

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 beetle.

Some species of beetle are kept as pets , for example diving beetles Dytiscidae may be kept in a domestic fresh water tank. In Japan the practice of keeping horned rhinoceros beetles Dynastinae and stag beetles Lucanidae is particularly popular amongst young boys.

Beetle collecting became extremely popular in the Victorian era. 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.

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.

Research published in sought to create a robotic camera backpack for beetles. 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.

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.

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.

You've probably seen the wide array of unique pooper scoopers that we've featured in the past, with some that even disinfect the area afterwards.

But none of them have anything on this incredible new dog poop picker upper robot. It's called the Beetl, and it finds, detects, and automatically scoops up poop throughout your yard.

It's like a little yard Roomba that'll clean up your yard, and is perfect for families with lots of dogs. Once it finds it, the robot rolls directly over it, and drops down a mechanical claw that picks up it and places it into a sealed internal container to be disposed of once it's done.

Bettl