Dermestids are 2-12-mm-long oval or elongate oval beetles with short, clubbed antennae. They usually have a distinct color pattern, and many are covered with fine setae or scales. Full-grown larvae are 4-12 mm long, brown and usually with long setae on the sclerites and posterior end; they are usually slow-moving. Their primitive habitat was probably feeding on decomposing animal matter, but, as the group evolved, feeding habits radiated to include other material. They are general scavengers on plant and animal material, including carrion, leather, furs, skins, museum specimens, wool and silk, and stored-food products. Larvae are capable ofdigesting keratin, which is a proteinaceous constituent ofwool and other animal materials. Some dermestids visit flowers, and several species require pollen for successful egg production. Adults are capable flyers and can move indoors through doors and windows. About 55 species of dermestids are primary or secondary pests of stored foods. Many dermestid pests made the transition to household habitats from living in insect and bird nests and animal burrows. Dermestid beetles are among the commonest inhabitants of bird nests (especially sparrows), and Anthrenus verbasci is commonly found in abandoned wasp nests, including those ofPolistes and Dolichovespula.

The family includes such well-knownpests as hide and larder beetles, Dermestes spp., and carpet beetles, Anthrenus spp. and Attagenus spp., which feed on materials of animal origin. This diverse family also includes the kaphra beetle, Trogoderma gran-arium, which feeds exclusively on plant material and is a major pest of grain and cereal products. The larval stage of dermestids damages household materials; they can have an extended period of development and move to several locations while feeding. Full-grown larvae pupate in the last larval skin (Anthrenus spp.), or it sheds the skin (Attagenus spp.). Dermestes larvae bore into solid material to construct a pupal chamber, and then the last larval skin and larval debris is used to block the entrance to the chamber. Development of the adult is often followed by a short resting period within the last larval skin. After mating, the female lays eggs on a suitable substrate and then usually becomes positively phototactic and may be found at windows and indoor lights.

Some species of carpet beetles are named for the plants on which the adult has been found feeding. Anthrenus scrophulariae is named for the figwort, Scrophularia spp.; A. verbasci is named for the mullein, Verbascum spp., and a common name for this plantis mothmullein, because itapparently attracts moths. The bird nestcarpet beetle A. pimpinellaeis named for the pimpernel, Anagallis spp. (= Pimpinella), presumably because adults occur on the flowers of this plant. Larvae of Attagneus feed on a variety of dry proteinaceous materials. The most common habitats for dermestid larvae are bird nests and rodent nests, but they also feed in bee and wasp nests and spider webs. Adults are often found on flowers feeding on nectar and pollen. Many species have adapted to living indoors and successive generations are produced withoutaccess to outdoors. The species thatare general feeders and tolerant of indoor conditions are household pests around the world.

Pest status of dermestid beetles is based primarily on the damage to commercial and household food materials, and the infestations that occur in the stored ingredients to the finished food products. Feeding on organic fabrics and other domestic materials results in economic loss and replacement costs. Infestations in museums result in losses to scientific collections ofinsects and vertebrates, and archived fabric and furs. Infestations may also cause medical problems. Setae on the larvae easily detach and produce allergic reactions, such as rhinitis and respiratory asthma. Infestations of Dermestes larvae indoors can expose sensitive skin and result in irritation. Similarly infested ship's cargo can release large quantities of larval skins in a confined space and cause irritated skin, conjunctivitis, and irritation of respiratory passages. The barbed larval setae of Trogoderma may occur in large numbers in infested grain, and cause allergic reactions if swallowed in food processed from this grain. D. maculatus and D. lardarius are pests in deep-pit poultry houses. The design of these commercial operations provides warm conditions and a readily available food supply for these beetles. Full-grown beetle larvae tunnel into woodwork and insulation in preparation for pupation, and these materials can become damaged and structurally

Carpet Beetle Skin Reaction
Figure 5.8 Coleoptera: Dermestidae adults. (a) Anthrenus flavipes; (b) A. fuscus; (c) A. museorum; (d) A. pimpinellae; (e) A. scrophularlae; (f) A. verbasci.

weakened. Infested manure spread on agricultural fields results in the infestation of nearby houses.

Asian carpet beetle, Anthrenus coloratus Adults are 1.5-2.5 mm long and mottled white and black. Antennal segments 4-6 are elongate, and the visible abdominal sternite 1 has pale white scales, while sternites 2-5 have yellow anterior scales and pale white posterior scales. This species closely resembles the varied carpet beetle, but is distinguished by nine antennal segments, whereas A. verbasci has 11 antennal segments. It is originally from Asia, Africa, and Europe, but recently it has been recorded from western North America. It occurs in the nests of wasps and spiders, and infesting insect collections, seeds, and various plant products.

Furniture carpet beetle, Anthrenus flavipes (= A. vorax) (Fig. 5.8a; 5.9i) Adults are 2-3.5 mm long. The body is rounded oval and spotted yellow, white, and black on the dorsum, and white ventrally. Full-grown larvae are about 5 mm long; they are widest at the anterior end and become narrow toward the posterior. Eggs are laid in 1-3 batches containing up to 57 eggs; fecundity is 37-96 eggs. Hatching occurs in 9-16 days; no eggs hatch at 40 °C and development is slow at 20 ° C. Adults overwinter, but no eggs are laid during cold weather. Larval development is 112-378 days depending on temperature; larvae that develop normally have six instars, but those developing slowly have 12 or more instars. Larvae feed in a limited radius and their cast larval skins can accumulate in one place, which gives the appearance of a severe infestation. The pupal period is 14-19 days, and the inactive period before emergence is 6-71 days. Development from egg to adult is 93-422 days, and adults live 30-60 days. Adults emerge in the spring and remain active during the warm season. Individuals that develop rapidly as larvae generally have a long adult life, and those that develop slowly have a short adult life.

This species is nearly cosmopolitan, but is probably native to the oriental region. It is named for its habit of attacking the animal-hair packing of stuffed furniture, which was a common practice in the early 1900s. It was first found in the USA in 1911 in upholstered furniture stuffed with horsehair imported from Russia. Larvae are sustained in modern households by feeding on a variety of organic substrates, including wool, silk, fur, feathers, and dry animal material. They need keratin in their diet and larvae will not fully develop if fed on pure wool unless it is impregnated with some other animal product. Larvae will feed on book bindings, chew holes in paper, and are attracted to and feed on dead animals.

Anthrenus fuscus (Fig. 5.8b) Adults are 1.7-2.8 mm long and with a variable pattern of black, white, and orange-brown scales on the elytra. It is distinguished from the varied carpet beetle, A. verbasci,byhaving five antennal segments, whereas A. verbasci has 11 antennal segments. Eggs are laid on dead insects in late spring, and larval development usually extends to the following year. Indoors it is found damaging woolens, carpets, insect collections, and books. Itis usually associated with spider webs where it feeds on insect debris that gathers in the web, or in corners where webs are built. Outdoors, larvae are found in peridomestic habitats, such as in outbuildings, sheds, barns, and stone walls. A./uscus usually does notoccur in bird nests, but is found associated with the sheet webs of the spider, Tegenaria domestica. This species occurs in North America, Europe, and Asia, including Japan.

Museum beetle, Anthrenus museorum (Fig. 5.8c) Adults are about 3 mm long and have yellow and white spots. It resembles A. verbasci,butthe antennae ofA. museorum are eight-segmented with a two-segmented club, whereas the antennae of A. verbasci are 11-segmented with a three-segmented club. Eggs are laid in late summer and larvae overwinter; pupation occurs in spring. Adults are active on flowers in sunlight, then become negatively phototactic and seek sheltered sites for oviposition.

There is usually one generation per year. This species is widely distributed, including North America, Europe, Australia, and New Zealand. In some regions, such as the UK, it lives primarily outdoors and is not a common indoor pest. Outdoors larvae are not usually found in bird nests, but have been found feeding on moth pupae in the nests of bees, and it occurs in the nests of spiders living in peridomestic habitats. In the sheet webs of T. domestica larvae are found in the egg sacs, feeding on eggs and spiders that failed to hatch. A. museorum larvae are capable of crawling on the sheet web of this spider, without being attacked. Pupation occurs among the remains ofinsects in the web or in old egg sacs. Indoors, this species attacks furs, woolens, carpets, silk, feathers, and skins. Larvae have also been recorded feeding on grain, wool, silk, and museum specimens, and dead cluster flies (Pollenia rudis) in attics.

Anthrenus olgae Adults are 1.7-2.8 mm long and there are two distinct pale white bands on the elytra. Antennae are eight-segmented and have a distinct club. The abdominal sternites are uniformly covered with pale white scales. Distribution of this species was originally limited to Europe, but it has been introduced into the UK. It is not a common indoor pest, and has been recorded from insect and bird collections.

Bird nest carpet beetle, panda carpet beetle, Anthrenus pimpinellae (= A. pimpinellas lepidus) (Fig. 5.8d) Adults are 2-4.5 mm long and are mottled brown and white. The prono-tum posterior margin has a patch of pale white scales at each side, which typically enclose a small oval, dark patch of scales. Full-grown larvae are about 5 mm long and blackish brown. Eggs are laid in the spring or early summer; fecundity is about 50 eggs; hatching occurs in 8 days at 26 °C and 15 days at 2022 °C. Larval development takes 3-4 months, and the pupal stage lasts 8-10 days; larvae pupate in the last larval skin. Adults develop in the fall but overwinter and become active in the spring. They feed on pollen and nectar. This species is nearly cosmopolitan, and occurs indoors and outdoors inmost regions of the world. In North America and Europe, it has been found damaging dried fish, woolens, and other animal materials. Outdoors, it develops in bird nests where the larvae feed on feathers, dead nestlings, or insect remains. Adults are common on flowers in the spring, then they move to bird nests or inside dwellings for oviposition.

Guernsey carpet beetle, Anthrenus sarnicus Adults are 2.6-3.2 mm long and have an indistinct mottled pattern. Antennae are 10-segmented, and segments 4-6 are rounded and beadlike. Abdominal sternites are uniformly covered with grayish-white scales. Eggs are laid in the spring, and fecundity is 50-80 eggs; hatching occurs in about 1.5 weeks. Larval development is completed in about 104 weeks at 15 °C, 40-50 weeks at 20 °C, and 10-18 weeks at 25 °C. The pupal period is 1.5-2.5 weeks. The adults live 3-14 weeks. This species was described from specimens collected in a house in Guernsey, Channel Isles, UK, and it has been found in domestic and commercial sites. It has been collected outdoors on flowers, and larvae have been recovered from house sparrow and pigeon nests, where it feeds on droppings, feathers, dead birds and insects. Larvae generally feed on animal materials, and indoors they infest fur, feather, skins, and dead insects.

Common carpet beetle, marbled carpet beetle, buffalo carpet beetle, Anthrenus scrophulariae (Fig. 5.8e; 5.9h) Adults are about 3 mm long. The body is oval, gray to black, and with a varied pattern of white and orange-red scales on the dorsal surface. The antennal club has segment 1 distinctly shorter than segment 2. Full-grown larvae are about 3 mm long, reddish brown, and with long setae. Eggs are laid singly or in batches of up to 36, and they are deposited directly on suitable food; fecundity is 40-60 eggs. Hatching occurs in 13-20 days, and the larval development period through about six instars is 60-80 days. First-stage larvae molt in about 13 days, second-, third-, and fourth-stage larvae molt in 8-10 days, while fifth-and sixth-stage larvae molt in 11-18 days. Pupae are formed in the last larval skin, and the pupal period is 7-12 days. The adult remains in the pupal chamber for 14-18 days, then it is active for about 30 days. Development from egg to adult takes about 95 days, and ranges from 89 to 108 days. Adults copulate and feed on the blossoms ofwhite flowers; the females require nectar and pollen to stimulate oviposition. This requirement usually limits indoor infestations, and links long-term household infestations to adults moving in and out of dwellings. Once mated and fed, females fly into houses or they are carried in on flowers such as daisies (Chrysanthemum), wild asters, white roses, crepe myrtle (Lagerstroemia indica), and shrubs such as lilac (Ceanothis)and Spiraea. Once inside, or in an animal burrow or nest, the female lays eggs and dies. Larvae feed until the end of the warm season, when about 75% pupate, with the remainder overwintering.

This is a cosmopolitan household pest, butmore common in north temperate regions, and less common in humid regions. In the UK it is only known from imported material and not known to occur outdoors; in other countries, such as Finland, it is only known to occur outdoors in bird and wasp nests and

Attagenus Brunneus Larva
Figure 5.9 Coleoptera: Dermestidae larvae. (a) Dermestes ater; (b) D. lardarius; (c) D. maculatus; (d) Trogoderma variabile; (e) T. versicolor; (f) Attagenus unicolor; (g) Anthrenus verbasci; (h) A. scrophulariae; (i) A.flavipes.

on dead animals. There seem to be two biological variants: one breeds indoors and does not visit flowers, and one breeds outdoors. Carpetbeetlelarvae feed on various animal materials, including wool, feathers, hair, and fur, museum specimens, and dried plants.

Varied carpet beetle, varied cabinet beetle, small cabinet beetle, Anthrenus verbasci (Fig. 5.8f 5.9g) Adults are 2-3 mm long. The dorsal surface of the body has a pattern of white, black, and brownish yellow scales, and the ventral surface has grayish-yellow scales. Full-grown larvae are 4-5 mm long and have a series of light- and dark-brown transverse stripes. Late-stage larvae are broad toward the rear and narrow at the front, and at each side on the posterior end of the body there are tufts of long and short setae. When alarmed the larva erects these tufts of setae and rolls into a ball as a defensive response. Eggs are deposited singly or in batches; fecundity is about 40 eggs. Hatching is in about 18 days; it is in 30-35 days at 18 °C, 4-17 days at 24 °C, and 10-12 days at 29 °C. Larval development takes 222-323 days, and includes 5-16 instars; the developmentperiod is determined by temperature, humidity, and food quality. Successful larval development is between 15-25 °C. Pupation is in the last larval skin. Adults remain inactive in the pupal case for 1-8 days, then emerge. The pupal period is 10-13 days; it is 17-19 days at 18 °C, 10-12 days at 24 °C, and 7-8 days at29 °C. Field strains of A. verbasci have an apparent diapause and must experience cold temperatures to produce adults in the spring; household strains produce adults in the fall. Diapause depends primarily on temperature. This species has one diapause at 25 °C and completes development in 1 year; there are two diapause stages at 15 °C, and the life cycle takes 2 years.

This species is cosmopolitan and occurs indoors and outdoors, but it is primarily a household pest on plant (dried fruits and nuts) and animal materials. A. verbasci is a pest of stored food materials and products, biscuits, cakes, seeds, wheat, maize, oats, rice, cayenne pepper, cacao, and dried cheese. It occurs in nests of birds, such as sparrows, martins, and swallows, and in wasp nests in attics, and under the siding of houses. It has also been found in bat roosts. In late spring and early summer, adults are found outdoors on as many as 30 species of flowering plants, but especially Spiraea, feeding on pollen. Adults from indoor populations have a negative attraction to light, but near the end of their oviposition period they become positively attracted to light. Adults from outdoor populations show a positive attraction to light. Adults are active flyers and often fly high above the ground. They enter houses through open windows, around eaves, soffits, and attic vents. This beetle is a common pest ofinsect collections, and the feeding larvae often leave a ring offrass around the hollowed-out insects. Adults have been reported to lay eggs in the dead insects collecting in light fixtures. Hymenoptera parasites recorded from A. verbasci include Laelius trogodermis and L. anthrenivorus (Bethylidae).

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