|Subfamilies, Genera & Species|
Bed bugs are parasitic insects of the cimicid family that feed exclusively on blood. The term most commonly refers to members of the genus Cimex of which Cimex lectularius, the common bed bug, is the best known as it prefers to feed on human blood although other Cimex species are specialized to other animals, e.g., bat bugs, Cimex pipistrelli (Europe), Cimex pilosellus (western US), and Cimex adjunctus (entire eastern US).
The name of the "bed bug" is derived from the preferred habitat of Cimex lectularius: warm houses and especially nearby or inside of beds and bedding or other sleep areas. Bed bugs are mainly active at night, but are not exclusively nocturnal. They usually feed on their hosts without being noticed.
A number of adverse health effects may result from bed bug bites, including skin rashes, psychological effects, and allergic symptoms. Diagnosis involves both finding bed bugs and the occurrence of compatible symptoms.
Bed bugs have been known as human parasites for thousands of years. At a point in the early 1940s, they were mostly eradicated in the developed world, but have increased in prevalence since 1995. Because infestation of human habitats has been on the increase, bed bug bites and related conditions have been on the rise as well.
- 1 Infestation
- 2 Description
- 3 Epidemiology
- 4 History
- 5 Society and culture
- 6 Further reading
- 7 References
- 8 External links
Bed bugs can cause a number of health effects, including skin rashes, psychological effects, and allergic symptoms. They are able to be infected by at least 28 human pathogens, but no study has clearly found that the insect is able to transmit the pathogen to a human being. Bed bug bites or cimicosis may lead to a range of skin manifestations from no visible effects to prominent blisters.
Diagnosis involves both finding bed bugs and the occurrence of compatible symptoms. Treatment involves the elimination of the insect and measure to help with the symptoms until they resolve. They have been found with methicillin-resistant Staphylococcus aureus (MRSA) and with vancomycin-resistant Enterococcus faecium (VRE), but the significance of this is still unknown.
Dwellings can become infested with bed bugs in a variety of ways, such as:
- Bugs and eggs inadvertently brought in from other infested dwellings on a visiting person's clothing or luggage;
- Infested items (such as furniture, clothing, or backpacks) brought in;
- Nearby dwellings or infested items, if easy routes are available for travel (through duct work or false ceilings);
- Wild animals (such as bats or birds) that may also harbour bed bugs or related species such as the bat bug;
- People visiting an infested area (apartment, subway, movie theatre, or hotel) and carrying the bugs to another area on their clothing, luggage, or bodies.
- Though bed bugs will feed on pets, they do not live or travel on the skin of their hosts, and pets are not believed to be a factor in their spread.
Bed bugs are elusive and usually nocturnal (peak activity usually occurs between 10:00 p.m. - 6:00 a.m.), which can make their detection difficult. They often lodge in dark crevices, and the tiny adhesive eggs can be nestled by the hundreds in fabric seams. Aside from bite symptoms, signs include fecal spots (small dark sand-like droppings that occur in patches around and especially beneath nests), blood smears on sheets (fecal spots that are re-wetted will smear like fresh blood), and the presence of their empty moulted exoskeletons.
Although bed bugs can be found singly, they tend to congregate once established. Although they are strictly parasitic, they spend only a tiny fraction of their life cycles physically attached to their hosts. Once feeding is complete, a bed bug will relocate to a place close to a known host, commonly in or near beds or couches in clusters of adults, juveniles, and eggs which entomologists call harborage areas or simply harborages to which the insect will return after future feedings by following chemical trails. These places can vary greatly in format, including luggage, inside of vehicles, within furniture, amongst bedside clutter, even inside electrical sockets and nearby laptop computers. Bed bugs may also nest near animals that have nested within a dwelling, such as bats, birds, or rodents. They are also capable of surviving on domestic cats and dogs, though humans are the preferred host of Cimex lectularius.
Bed bugs can also be detected by their characteristic smell of rotting raspberries. Bed bug detection dogs are trained to pinpoint infestations, with a possible accuracy rate of between 11 and 83%. A few companies are experimenting with high speed gas chromatography to detect bed bugs.
Mechanical approaches, such as vacuuming up the insects and heat treating or wrapping mattresses, are effective. A combination of heat and drying treatments has been found to be most effective. An hour of 45 degree C temperature will kill them. Temperatures of less than -17 degrees C for 2 hours are also effective. Starving them is difficult as they can survive without eating for 100 to 300 days depending on the temperature. For public health reasons, individuals are encouraged to call a professional pest control service to eradicate bed bugs in a home, rather than attempting to do it themselves, particularly if they live in a multi-family building.
As of 2012 there are no really effective pesticides. Pesticides that have historically been found to be effective include: pyrethroids, dichlorvos and malathion. Resistance to pesticides has increased significantly over time and negative health effects from their use are of concern. The carbamate insecticide propoxur is highly toxic to bed bugs, but in the United States, the Environmental Protection Agency (EPA) has been reluctant to approve it for indoor use because of its potential toxicity to children with exposure. Although occasionally applied as a safe indoor pesticide treatment for other insects, boric acid is not effective against bed bugs because bed bugs do not groom. The fungus Beauveria bassiana is being researched for its ability to control bed bugs.
Natural enemies of bed bugs include the masked hunter insect (also known as "masked bed bug hunter"), cockroaches, ants, spiders (particularly Thanatus flavidus), mites and centipedes (particularly the house centipede Scutigera coleoptrata). Biological pest control is currently not considered practical for eliminating bed bugs from human dwellings.
Adult bed bugs are light brown to reddish-brown, flattened, oval-shaped and have no hind wings. The front wings are vestigial and reduced to pad-like structures. Bed bugs have segmented abdomens with microscopic hairs that give them a banded appearance. Adults grow to 4–5 mm in length and 1.5–3 mm wide.
Newly hatched nymphs are translucent, lighter in color and become browner as they moult and reach maturity. A bed bug of any age that has just consumed a blood meal will appear to have a bright red translucent abdomen; this color will fade to brown over the next several hours and within two days will become opaque and black as the insect digests its meal. Bed bugs may be mistaken for other insects, such as booklice, small cockroaches, or carpet beetles, however when warm and active, their movements are more ant-like, and like most other true bugs, they emit a characteristic disagreeable odor when crushed.
The life span of bed bugs varies by species and is also dependent on feeding.
Bed bugs can survive a wide range of temperatures and atmospheric compositions. Below 16.1 °C (61.0 °F), adults enter semihibernation and can survive longer; they can survive for at least five days at −10 °C (14 °F), but will die after 15 minutes of exposure to −32 °C (−26 °F). They show high desiccation tolerance, surviving low humidity and a 35–40 °C range even with loss of one-third of body weight; earlier life stages are more susceptible to drying out than later ones.
The thermal death point for Cimex lectularius is high: 45 °C (113 °F), and all stages of life are killed by 7 minutes of exposure to 46 °C (115 °F). Bed bugs apparently cannot survive high concentrations of carbon dioxide for very long; exposure to nearly pure nitrogen atmospheres, however, appears to have relatively little effect even after 72 hours.
Bed bugs are obligatory hematophagous (bloodsucking) insects. Most species feed on humans only when other prey are unavailable. They obtain all the additional moisture they need from water vapor in the surrounding air. Bed bugs are attracted to their hosts primarily by carbon dioxide, secondarily by warmth, and also by certain chemicals. Bedbugs prefer exposed skin, preferably the face, neck and arms of a sleeping individual.
Although under certain cool conditions adult bed bugs can live for over a year without feeding, under typically warm conditions they will try to feed at five- to ten-day intervals and adults can survive for about five months without food. Younger instars cannot survive nearly as long, though even the vulnerable newly hatched first instars can survive for weeks without taking a blood meal.
A bed bug pierces the skin of its host with what is called a stylet fascicle, rostrum, or "beak". The rostrum is composed of the maxillae and mandibles, which have been modified into elongated shapes from a basic, ancestral style. The right and left maxillary stylets are connected at their midline and a section at the centerline forms a large food canal and a smaller salivary canal. The entire maxillary and mandibular bundle penetrates the skin.
The tips of the right and left maxillary stylets are not the same; the right is hook-like and curved, and the left is straight. The right and left mandibular stylets extend along the outer sides of their respective maxillary stylets and do not reach anywhere near the tip of the fused maxillary stylets. The stylets are retained in a groove in the labium, and during feeding, they are freed from the groove as the jointed labium is bent or folded out of the way; its tip never enters the wound.
The mandibular stylet tips have small teeth and through alternately moving these stylets back and forth, the insect cuts a path through tissue for the maxillary bundle to reach an appropriately sized blood vessel. Pressure from the blood vessel itself fills the insect with blood in three to five minutes. The bug then withdraws the stylet bundle from the feeding position and retracts it back into the labial groove, folds the entire unit back under the head, and returns to its hiding place. It takes between five and ten minutes for a bed bug to become completely engorged with blood. In all, the insect may spend less than 20 minutes in physical contact with its host, and it will not attempt to feed again until it has either completed a moult or, if an adult, has thoroughly digested the meal.
All bed bugs mate by traumatic insemination. Female bed bugs possess a reproductive tract that functions during oviposition, but the male does not use this tract for sperm insemination. Instead, the male pierces the female's abdomen with his hypodermic genitalia and ejaculates into the body cavity. In all bed bug species except Primicimex cavernis, sperm are injected into the mesospermalege, a component of the spermalege, a secondary genital structure that reduces the wounding and immunological costs of traumatic insemination. Injected sperm travel via the haemolymph (blood) to sperm storage structures called seminal conceptacles, with fertilisation eventually taking place at the ovaries.
Bed bugs have six life stages (five immature nymph stages and a final sexually mature adult stage). They shed their skins through ecdysis at each stage, discarding their outer shells which are clear, empty exoskeletons of the bugs themselves. Bed bugs must molt six times before becoming fertile adults and must take a blood meal in order to complete each moult.
Each of the immature stages lasts approximately a week, depending on temperature and the availability of food, and the complete life cycle can be completed in as little as two months (which is actually rather long compared to other ectoparasites). Fertilized females with enough food will lay three to four eggs each day continually until the end of their life spans (about nine months under warm conditions), possibly generating as many as 500 eggs in this time.
Sexual dimorphism occurs in Cimex lectularius with the males larger in size than the females on average. The abdomens of the sexes differ in that the males have pointed abdomens, which hold the copulatory organ. Despite the physiological differences, males are unable to discriminate between the sexes until after mounting, but before inseminating.
Cimex lectularius only feed every five to seven days, which suggests that they do not spend the majority of their life searching for a host. When a bed bug is starved it will leave its shelter and search for a host. If it succeeds at finding food, it will return to its shelter. If it does not feed, it will continue to search for a host. After searching, regardless of whether or not it has eaten, the bed bug will return to the shelter to aggregate before the photophase (period of light during a day-night cycle). Reis argues that there are two reason why the Cimex lectularius would return to their shelter and aggregate after feeding. One of the reasons is to find a mate and the other is to find shelter to avoid getting smashed after eating.
Aggregation and dispersal behavior
Cimex lectularius aggregates under all life stages and mating conditions. Bed bugs may choose to aggregate because of predation, resistance to desiccation, and more opportunities to find a mate. Airborne pheromones are responsible for aggregations. Another source of aggregation could be the recognition of other Cimex lectularius through mechanoreceptors located on the antennae. Aggregations are formed and disbanded based on the cost and benefits associated. Females are more often found separate from the aggregation than males. Females are more likely to expand the population range and find new sites. Active female dispersal can account for treatment failures. Males, when found in areas with few females, will abandon an aggregation in order to find a new mate. The males excrete an aggregation pheromone into the air that attracts virgin females and arrests other males.
Male bed bugs sometimes attempt to mate with other males and pierce their abdomens. This behaviour occurs because sexual attraction in bed bugs is based primarily on size, and males will mount any freshly fed partner regardless of sex. The "bed bug alarm pheromone" consists of (E)-2-octenal and (E)-2-hexenal. It is released when a bed bug is disturbed, as during an attack by a predator. A 2009 study demonstrated the alarm pheromone is also released by male bed bugs to repel other males who attempt to mate with them.
Cimex lectularius and Cimex hemipterus will mate with each other given the opportunity, but the eggs then produced are usually sterile. In a 1988 study, one of 479 eggs was fertile and resulted in a hybrid, Cimex hemipterus × lectularius.
Cimex lectularius males are shown to have environmental microbes on their genitals. These microbes damage sperm cells leaving them unable to fertilize female gametes. Due to these dangerous microbes, male Cimex lectularius have evolved antimicrobial ejaculate substances which act to prevent sperm damage. Upon contact with the microbes with sperm or the male genitals, antimicrobial substances will be released. Many species of these microbes can also be found in the bodies of females after mating. The microbes can cause infections in the females. It has been suggested that females receive benefit from the ejaculate. Although the benefit is not direct, the females are able to produce more eggs than optimum increasing the amount of the females’ genes in the gene pool.
Sperm and seminal fluid allocation
In organisms, sexual selection extends past differential reproduction to affect sperm composition, sperm competition, and ejaculate size. It has been demonstrated that male C imexlectularius allocate 12% of their sperm and 19% of their seminal fluid per mating. Due to these findings, Reinhard et. al proposed that multiple mating is limited by seminal fluid and not sperm. After measuring ejaculate volume, mating rate and estimating sperm density, Reinhard et al. showed that mating could be limited by seminal fluid. Despite these advances, the cost difference between ejaculate-dose dependence and mating frequency dependence have not been explored.
Females are fertilized in two ways. The first is through vaginal intercourse and the second through forced copulation where the male forces himself into the organ called the Ribaga, which is to the right of the fourth apparent sternum. Upon fertilization, the female’s ovaries finish developing which suggests that sperm play a role other than fertilizing the egg. Fertilization also allows for egg production through the corpus allatum. Sperm remains viable in a female’s spermathecae, a sperm carrying sack, for a long period of time as long as body temperature is optimum. The females lays fertilized eggs until she depletes the sperm found in her spermathecae. After the depletion of sperm, she lays a few sterile eggs. The number of eggs a Cimex lectularius females produces does not depend on the sperm she produces but on the female’s nutritional level.
In Cimex lectularius, males sometimes mount other males. This is because male sexual interest is directed at any recently fed individual regardless of their sex. Traumatic insemination is the only way for copulation to occur in bed bugs. Females have evolved the spermalege to protect themselves from wounding and infection. Because males lack this organ, traumatic insemination could leave them injured badly. For this reason, males have evolved alarm pheromones to signal their sex to other males. If a male Cimex lectularius is mounted by another male, the mounted male will release the pheromone signal and the male on top will know to stop before insemination. Females are able to produce alarm pheromones to avoid multiple mating but they do not do so. There are two proposed reasons why females do not release alarm pheromones to protect themselves. First, alarm pheromone production is costly. It has been suggested that due to egg production, females refrain from spending additional energy on alarm pheromones. The second proposed reason is that releasing the alarm pheromone reduces the benefits associated with multiple mating. Benefits of multiple mating include material benefits, better quality nourishment or more nourishment, genetic benefits including increased fitness of offspring, and finally, which appears to be the case in Cimex lectularius, the cost of resistance may be higher than the benefit of consent.
Bed bugs occur around the world. Rates of infestations in developed countries, while decreasing from the 1930s to the 1980s, have increased dramatically since the 1980s. Previously, they were common in the developing world, but rare in the developed world. The increase in the developed world may have been caused by increased international travel, resistance to insecticides, and the use of new pest-control methods that do not affect bed bugs.
The fall in bed bug populations after the 1930s in the developed world is believed to be partly due to the use of DDT to kill cockroaches. The invention of the vacuum cleaner and simplification of furniture design may have also played a role. Others believe it might simply be the cyclical nature of the organism.
The exact causes of this resurgence remain unclear; it is variously ascribed to greater foreign travel, increased immigration from the developing world to the developed world, more frequent exchange of second-hand furnishings among homes, a greater focus on control of other pests, resulting in neglect of bed bug countermeasures, and increasing resistance to pesticides.
The common bed bug (Cimex lectularius) is the species best adapted to human environments. It is found in temperate climates throughout the world. Other species include Cimex hemipterus, found in tropical regions, which also infests poultry and bats, and Leptocimex boueti, found in the tropics of West Africa and South America, which infests bats and humans. Cimex pilosellus and Cimex pipistrella primarily infest bats, while Haematosiphon inodora, a species of North America, primarily infests poultry.
Bed bugs were mentioned in ancient Greece as early as 400 BC, and were later mentioned by Aristotle. Pliny's Natural History, first published circa 77 AD in Rome, claimed bed bugs had medicinal value in treating ailments such as snake bites and ear infections. (Belief in the medicinal use of bed bugs persisted until at least the 18th century, when Guettard recommended their use in the treatment of hysteria.)
Bed bugs were first mentioned in Germany in the 11th century, in France in the 13th century and in England in 1583, though they remained rare in England until 1670. Some in the 18th century believed bed bugs had been brought to London with supplies of wood to rebuild the city after the Great Fire of London (1666). Giovanni Antonio Scopoli noted their presence in Carniola (roughly equivalent to present-day Slovenia) in the 18th century.
Traditional methods of repelling and/or killing bed bugs include the use of plants, fungi, and insects (or their extracts), such as black pepper; black cohosh (Actaea racemosa); Pseudarthria hookeri; Laggera alata (Chinese yángmáo cǎo | 羊毛草); Eucalyptus saligna oil; henna (Lawsonia inermis or camphire); "infused oil of Melolontha vulgaris" (presumably cockchafer); fly agaric (Amanita muscaria); Actaea spp. (e.g. black cohosh); tobacco; "heated oil of Terebinthina" (i.e. true turpentine); wild mint (Mentha arvensis); narrow-leaved pepperwort (Lepidium ruderale); Myrica spp. (e.g. bayberry); Robert geranium (Geranium robertianum); bugbane (Cimicifuga spp.); "herb and seeds of Cannabis"; "opulus" berries (possibly maple or European cranberrybush); masked hunter bugs (Reduvius personatus), "and many others".
Dusts have been used to ward off insects from grain storage for centuries, including "plant ash, lime, dolomite, certain types of soil, and diatomaceous earth or Kieselguhr". Of these, diatomaceous earth in particular has seen a revival as a nontoxic (when in amorphous form) residual pesticide for bed bug abatement. Insects exposed to diatomaceous earth may take several days to die.
Basket-work panels were put around beds and shaken out in the morning in the UK and in France in the 19th century. Scattering leaves of plants with microscopic hooked hairs around a bed at night, then sweeping them up in the morning and burning them, was a technique reportedly used in Southern Rhodesia and in the Balkans.
Bean leaves have been used historically to trap bedbugs in houses in Eastern Europe. The trichomes on the bean leaves capture the insects by impaling the feet (tarsi) of the insects. The leaves are then destroyed.
Prior to the mid-20th century, bed bugs were very common. According to a report by the UK Ministry of Health, in 1933, all the houses in many areas had some degree of bed bug infestation. The increase in bed bug populations in the early 20th century has been blamed on the advent of electric heating, which allowed bed bugs to thrive year-round instead of only in warm weather.
Bed bugs were a serious problem at U.S. military bases during World War II. Initially, the problem was solved by fumigation, using Zyklon Discoids that released hydrogen cyanide gas, a rather dangerous procedure. Later, DDT was used to good effect as a safer alternative.
The decline of bed bug populations in the 20th century is often credited to potent pesticides that had not previously been widely available. Other contributing factors that are less frequently mentioned in news reports are increased public awareness and slum clearance programs that combined pesticide use with steam disinfection, relocation of slum dwellers to new housing, and in some cases also follow-up inspections for several months after relocated tenants moved into their new housing.
Bed bug infestations have resurged in recent years for reasons that are not clear, but contributing factors may be complacency, increased resistance, bans on pesticides and increased international travel. The U.S. National Pest Management Association reported a 71% increase in bed bug calls between 2000 and 2005. The number of reported incidents in New York City alone rose from 500 in 2004 to 10,000 in 2009.
One recent theory about bed bug reappearance is that they never truly disappeared from the United States, but may have been forced to alternative hosts. Consistent with this is the finding that bed bug DNA shows no evidence of an evolutionary bottleneck. Furthermore, investigators have found high populations of bed bugs at poultry facilities in Arkansas. Poultry workers at these facilities may be spreading bed bugs, unknowingly carrying them to their places of residence and elsewhere after leaving work.
Society and culture
The saying, "Good night, sleep tight, don't let the bed bugs bite," is common for parents to say to young children before they go to sleep. In Chhattisgarh, India, bed bugs, have been used as a traditional medicine for epilepsy, piles, alopecia and urinary disorders; however this practice has no scientific basis. Bed bug saliva however is suspected of having antibacterial proteins which could one day be used to create new drugs.
The word bug and its earlier spelling bugge means bedbug. Many other creatures are called bugs, such as the lady bug (ladybird outside North America), the potato bug, or the informal use of the word for any insect, or even for microscopic germs, or diseases caused by these germs, but the earliest recorded use of the actual word bug was to mean bedbug.
The term bed bug may also be spelled bedbug or bed-bug, though published sources consistently use the unhyphenated two-word name bed bug. They have been known by a variety of other names, including wall louse, mahogany flat, crimson rambler, chilly billies, heavy dragoon, chinche bug, and redcoat.
- Stephen Doggett. Bed Bugs: Clinical Relevance and Control Options. Clinical Microbiological Reviews, 25(1):164–192.
- Stephen Doggett. A Code of Practice for the Control of Bed Bugs in Australia. Draft 4th edition, ICPMR & AEPMA, Sydney Australia, September 2011. ISBN 1-74080-135-0.
- Stephen Doggett. A Bed Bug Management Policy for Accommodation Providers. First ed, ICPMR, Sydney Australia, Sep 2011.
- David Cain, Richard Strand. Bed Bug Beware: An easy to understand guide to bed bugs, their prevention and control. Loughborough, United Kingdom: Foxhill Publishing, March 2009. ISBN 978-0-9562617-0-0
- Larry Pinto, Richard Cooper, Sandy Kraft. Bed Bug Handbook: The Complete Guide to Bed Bugs and Their Control. Mechanicsville, Maryland: Pinto & Associates, December 2007. ISBN 978-0-9788878-1-0
- Forsyth, Adrian. A Natural History of Sex: The Ecology and Evolution of Mating Behavior. Richmond Hill, Ontario: Firefly Books, 2001. ISBN 1-55209-481-2.
- MacQuitty, Miranda, and Lawrence Mound. Megabugs: The Natural History Museum Book of Insects. New York: Random House Children's Books, 1995. ISBN 1-898304-37-8, ISBN 1-85868-045-X.
- Goddard, Jerome A. The Physician's Guide to Arthropods of Medical Importance (second edition).
- [No authors listed] (1996). "Oeciacus hirundinis (Lamarck, 1816)". Integrated Taxonomic Information System. Retrieved 2010-09-15.
- Bat bugs are visually indistinguishable from bed bugs, and may also occasionally feed on human hosts.
- Goddard, J; deShazo, R (1 April 2009). "Bed bugs (Cimex lectularius) and clinical consequences of their bites". JAMA: the Journal of the American Medical Association 301 (13): 1358–66. doi:10.1001/jama.2009.405. PMID 19336711.
- Reinhardt, Klaus; Siva-Jothy, Michael T. (Jan 2007). "Biology of the Bed Bugs (Cimicidae)". Annual Review of Entomology 52: 351–374. doi:10.1146/annurev.ento.52.040306.133913. PMID 16968204. Archived from the original on 5 July 2010. Retrieved 26 May 2010.
- "What Are Bed Bugs? How To Kill Bed Bugs". Medical News Today. MediLexicon International Ltd. 20 July 2009. Retrieved 27 May 2010.
- Doggett SL, Russell R (November 2009). "Bed bugs – What the GP needs to know". Aust Fam Physician 38 (11): 880–4. PMID 19893834.
- Jerome Goddard & Richard deShazo (2009). "Bed bugs (Cimex lectularius) and clinical consequences of their bites". Journal of the American Medical Association 301 (13): 1358–1366. doi:10.1001/jama.2009.405. PMID 19336711.
- Kilpenen, O.; Vagn Jensen, K-M.; Kristensen, M. Bed Bug Problems in Denmark, with a European Perspective. pp. 395–399. Retrieved 27 May 2010.
- Robinson, WH; Bajomi, D. (eds) (2008). Proceedings of the Sixth International Conference on Urban Pests. Hungary: OOK-Press Kft.
- Kolb A, Needham GR, Neyman KM, High WA (2009). "Bedbugs". Dermatol Ther 22 (4): 347–52. doi:10.1111/j.1529-8019.2009.01246.x. PMID 19580578.
- Kolb, A., et al. "Bedbugs." Dermatologic Therapy 22.4 (2009): 347–352. CAB Abstracts. EBSCO. Web. 5 December 2010.
- James, William D.; Berger, Timothy G. (2006). Andrews' Diseases of the Skin: clinical Dermatology. Saunders Elsevier. ISBN 0-7216-2921-0.
- Melnick, Meredith (2011-05-12). "Study: Bedbugs May Carry MRSA; Germ Transmission Unclear | TIME.com". Healthland.time.com. Retrieved 2013-11-11.
- "Do Bedbugs Spread MRSA and VRE?". Webmd.com. 2011-05-11. Retrieved 2013-11-11.
- Potter, Michael F. "BED BUGS". University of Kentucky College of Agriculture. Archived from the original on 2 July 2010. Retrieved 27 June2010.
- Steelman, C.D. 2000. Biology and control of bed bugs, Cimex lectularius, in poultry houses. Avian Advice 2: 10,15. 
- "The Truth About Bedbugs: Debunking the Myths". PAWS SF. Retrieved 13 Oct 2013.
- Susan L. Woodward; Joyce A. Quinn (30 September 2011). Encyclopedia of Invasive Species: From Africanized Honey Bees to Zebra Mussels: From Africanized Honey Bees to Zebra Mussels. ABC-CLIO. p. 124. ISBN 978-0-313-38221-5. Retrieved 15 August 2013.
- Anderson, AL; Leffler, K (May 2008). "Bedbug infestations in the news: a picture of an emerging public health problem in the United States". Journal of environmental health 70 (9): 24–7, 52–3. PMID 18517150.
- Doggett, SL; Dwyer, DE; Peñas, PF; Russell, RC (2012 Jan). "Bed bugs: clinical relevance and control options.". Clinical microbiology reviews 25 (1): 164–92. PMID 22232375.
- 2012, Scientific American, Hayes
- York Times. In Search of a Bedbug Solution. Published: 4 September 2010.
- Miller, Dini (2008). "Bed bugs (hemiptera: cimicidae: Cimex spp.)". In Capinera, John L. Encyclopedia of Entomology (Second ed.). Springer. p. 414. ISBN 978-1-4020-6242-1.
- "Natural Fungus May Provide Effective Bed Bug Control".
- Hahn, Jeffrey (November 1999). "Masked hunters". Yard & Garden Briefs. University of Minnesota Extension Service: Yard & Garden Clinic. Retrieved 27 May 2010.
- Encyclopedia Americana, 1996 ed., v. 3, p. 413
- "Bed Bugs Bounce Back". Birc.org. Retrieved 2013-11-11.
- Benoit, J. B.; del Grosso, N.; Yoder, J. A.; Denlinger, D. L. (2007). "Resistance to dehydration between bouts of blood feeding in the bed bug, Cimex lectularius, is enhanced by water conservation, aggregation, and quiescence". American Journal of Tropical Medicine and Hygiene 76 (5): 987–993. ISSN 0002-9637. PMID 17488928. Retrieved 27 May 2010.
- Herrmann, J.; Adler, C.; Hoffmann, G.; Reichmuth, C. (1999). "Efficacy of controlled atmospheres on Cimex lectularius (L.) (Heteroptera: Cimicidae) and Argas reflexus Fab. (Acari: Argasidae)". In Robinson, Wm H.; Rettich, F.; Rambo, G.W. Proceedings of the 3rd International Conference on Urban Pests. Hronov, Czech Republic: Grafické Závody. p. 637. Retrieved 31 May 2010. (abstracted from a poster presentation in Prague, 19–22 Jul)
- Storey, Malcom. "CIMICIDAE (bed bugs)". BioImages: The Virtual Field-Guide (UK). bioimages.org.uk. Retrieved 27 May 2010.
- Lance A., Gary R.; Durden (8 May 2009). Medical and Veterinary Entomology, Second Edition. Academic Press. p. 80. ISBN 0-12-372500-3.
- "Family CIMICIDAE". Australian Biological Resources Study: Australian Faunal Directory. Department of the Environment, Water, Heritage and the Arts (Australia). 2008. Retrieved 27 May 2010.
- Richards, Luck (1999). Integument of Arthropods. University of Minnesota. p. 298. ISBN 978-0-8166-0073-1.
- Anderson, J. F.; Ferrandino, F. J.; McKnight, S.; Nolen, J.; Miller, J. (2009). "A carbon dioxide, heat and chemical lure trap for the bed bug, Cimex lectularius". Medical and Veterinary Entomology 23 (2): 99–105. doi:10.1111/j.1365-2915.2008.00790.x. Retrieved 2010-05-27.
- Narinderpal Singh; Changlu Wang; Richard Cooper; Chaofeng Liu. "Interactions among Carbon Dioxide, Heat, and Chemical Lures in Attracting the Bed Bug, Cimex lectularius L. (Hemiptera: Cimicidae)". Psyche 2012. doi:10.1155/2012/273613.
- Changlu Wang; Gary W Bennett (2009). "Bed bug (Heteroptera: Cimicidae) attraction to pitfall traps baited with carbon dioxide, heat, and chemical lure.". Journal of Economic Entomology (8). 102(4):1580-5.
- Robert L. Usinger (1966). Monograph of Cimicidae (Hemiptera - Heteroptera) V11. The Thomas Say Foundation. p. 13.
- Polanco, Andrea (2011). "Survivorship During Starvation for Cimex lectularius L.". Insects 2: 232–242. doi:10.3390/insects2020232.
- Milius, Susan (16 January 2010). "Do-it-yourself bed-bug detector". Science News (Society for Science and the Public) 177 (2): 88. Retrieved 27 May 2010. citingPolanco-Pinzón, Andrea; et al. (2009). "Survivorship and growth potential of modern bed bug populations (Cimex lectularius) in the United States". Conference proceedings of 57th Annual Meeting (Entomological Society of America).
- A. L. Szalanski; J. W. Austin; J. A. McKern; C. D. Steelman; D. Miller (2006). "Time course analysis of bed bug, Cimex lectularius L., (Hemiptera: Cimicidae) blood meals using PCR". Journal of Agricultural and Urban Entomology 23: 237–241.
- A. L. Szalanski, J. W. Austin, J. A. McKern, D. Miller, C. D. Steelman and R. E. Gold. 2006. Isolation and characterization of human DNA from the bed bug, Cimex lectularius L. (Heteroptera: Cimicidae). Journal of Agricultural and Urban Entomology 23: 189–194. 
- I. Thomas; G. G. Kihiczak; R. A. Schwartz (2004). "Bed bug bites: a review". International Journal of Dermatology 43 (6): 430–433. doi:10.1111/j.1365-4632.2004.02115.x. PMID 15186224.
- Carayon, J. 1959 Insémination par "spermalège" et cordon conducteur de spermatozoids chez Stricticimex brevispinosus Usinger (Heteroptera, Cimicidae). Rev. Zool. Bot. Afr. 60, 81–104.
- Morrow E. H., Arnqvist G. (2003). "Costly traumatic insemination and a female counter-adaptation in bed bugs". Proceedings of the Royal Society B 270 (1531): 2377–2381. doi:10.1098/rspb.2003.2514. PMC 1691516. PMID 14667354.
- Reinhardt K., Naylor R., Siva-Jothy M. T. (2003). "Reducing a cost of traumatic insemination: female bedbugs evolve a unique organ". Proceedings of the Royal Society B 270 (1531): 2371–2375. doi:10.1098/rspb.2003.2515. PMC 1691512. PMID 14667353.
- Ryne, C. In press. "Homosexual interactions in bed bugs: Alarm pheromones as male recognition signals." Animal Behaviour.doi:10.1016/j.anbehav.2009.09.033
-  p. 136
- Shukla; Upadhyaya (2009). Economic Zoology (Fourth ed.). Rastogi. p. 73. ISBN 978-81-7133-876-4.
- Domingue, Michael J., Matthew Kramer, and Mark F. Feldlaufer. "Sexual Dimorphism of Arrestment and Gregariousness in the Bed Bug (Cimex Lectularius) in Response to Cuticular Extracts from Nymphal Exuviae." Physiological Entomology 35.3 (2010): 203-13. Print.
- Reis, Matthew D., and Dini M. Miller. "Host Searching and Aggregation Activity of Recently Fed and Unfed Bed Bugs (Cimex Lectularius L.)." Insects 2.2 (2011): 186-94. Print.
- Pfiester, Margie, Philip G. Koehler, and Roberto M. Pereira. "Effect of Population Structure and Size on Aggregation Behavior Of(Hemiptera: Cimicidae)." Journal of Medical Entomology 46.5 (2009): 1015-020. Print
- "This Bedbug's Life", The New York Times, 7 August 2010.
- Harari A., Brockman H. J., Landholt P. J. (2000). "Intrasexual mounting in the beetle Diaprepes abbreviatus (L.)". Proceedings of the Royal Society B 267 (1457): 2071–2079. doi:10.1098/rspb.2000.1251. PMC 1690776. PMID 11416911.
- "Scent of alarm identifies male bed bugs". Science News. 2009-10-29. Retrieved 2013-11-11.
- Newberry, K. (July 1988). "Production of a hybrid between the bedbugs Cimex hemipterus and Cimex lectularius". Medical and Veterinary Entomology (The Royal Entomological Society) 2 (3): 297–300. doi:10.1111/j.1365-2915.1988.tb00199.x. PMID 2980186.
- Walpole, Debra E.; Newberry, K. (July 1988). "A field study of mating between two species of bedbug in northern KwaZulu, South Africa". Medical and Veterinary Entomology (The Royal Entomological Society) 2 (3): 293–296. doi:10.1111/j.1365-2915.1988.tb00198.x. PMID 2980185.
- Otti, Oliver, Klaus Reinhard, and Aimee P. McTighe. "In Vitro Antimicrobial Sperm Protection by an Ejaculate-like Substance." Functional Ecology 27.1 (2013): 219-26. Print.
- Reinhart, Klaus, Richard Naylor, and Michael T. Siva-Toth. "Male Mating Rate Is Constrained by Seminal Fluid Availability in Bedbugs, Cimex Lectularius." PLOS ONE 6.7 (2011): 1-8. PLOS ONE. Web
- Mellanby, Kenneth. "Fertilization and Egg Production in the Bed-bug, Cimex Lectularius L."Parasitology 31.02 (1939): 193. Print.
- Ryne, Camille. "Homosexual Interactions in Bed Bugs: Alarm Pheromones as Male Recognition Signals." Animal Behaviour 78.6 (2009): 1471-475. Print.
- Davies, Nicholas B.; Krebs, John R.; West, Stuart A. (2012-02-17). An Introduction to Behavioural Ecology (Kindle Locations 5432-5434). Wiley. Kindle Edition.
- Heukelbach, J; Hengge, UR (2009). "Bed bugs, leeches and hookworm larvae in the skin". Clinics in dermatology 27 (3): 285–90. doi:10.1016/j.clindermatol.2008.10.008. PMID 19362691 pmid 19362691
- A Romero, MF Potter, DA Potter, KF Haynes (2007). "Insecticide Resistance in the Bed Bug: A Factor in the Pest's Sudden Resurgence?". Journal of medical entomology 22 (2): 175–178. doi:10.1603/0022-2585(2007)44[175:IRITBB]2.0.CO;2. ISSN 0022-2585. Retrieved 2010-09-17.
- Owen, James (13 May 2004). "Bloodthirsty Bedbugs Stage Comeback in U.S., Europe". National Geographic News. National Geographic. Retrieved 31 May 2010.
- Krause-Parello CA, Sciscione P (April 2009). "Bedbugs: an equal opportunist and cosmopolitan creature". J Sch Nurs 25 (2): 126–32. doi:10.1177/1059840509331438. PMID 19233933.
-  p. 131
- Cranshaw, W.S.; Camper, M.; Peairs, F.B. (Feb 2009). "Bat Bugs and Bed Bugs". Colorado State University Extension. Retrieved 27 May 2010.
- "A dictionary of Greek and Roman antiquities - Sir William Smith - Google Boeken". Books.google.com. Retrieved 2013-11-11.
- John Southall. ""That soon after the Fire of London, in some of the new-built Houses they were observ'd to appear, and were never noted to have been seen in the old, tho' they were then so few, as to be little taken notice of; yet as they were only seen in Firr-Timber, 'twas conjectured they were then first brought to England in them; of which most of the new Houses were partly built, instead of the good Oak destroy'd in the old."". A Treatise of Buggs [sic], pp. 16–17.
- Johann Friedrich Wolff and Johann Philip Wolff. ""According to Scopoli's 2nd work (loc. cit.), found in Carniola and adjoining regions. According to Linnaeus' second work on exotic insects (loc. cit.), before the era of health, already in Europe, seldom observed in England before 1670."". Icones Cimicum descriptionibus illustratae, fourth fascicle (1804), p. 127.
- George Orwell, Down and Out in Paris and London, 1933
- Schaefer, C.W.; Pazzini, A.R. (28 July 2000). Heteroptera of Economic Importance. Boca Raton, FL: CRC Press. p. 525. ISBN 0-8493-0695-7.
- Kambu, Kabangu; Di Phanzu, N.; Coune, Claude; Wauters, Jean-Noël; Angenot, Luc (1982). "Contribution à l'étude des propriétés insecticides et chimiques d'Eucalyptus saligna du Zaïre (Contribution to the study of insecticide and chemical properties of Eucalyptus saligna from Zaire ( Congo))". Plantes Médicinales et Phytothérapie (Paris: Jouve) 16 (1): 34–38. hdl:2268/14438.
- "Getting Rid of Bed-Bugs". Grubstreet.rictornorton.co.uk. Retrieved 2013-11-11.
- "Icones Cimicum descriptionibus illustratae". Archive.org. Retrieved 2013-11-11.
- (no byline) (17 June 1848). "Peat and peat mosses". Scientific American 3 (39): 307. Retrieved 26 May 2010.
- Hill, Stuart B. (May 1986). "Diatomaceous Earth: A Non Toxic Pesticide". Macdonald J. (Ste-Anne de Bellevue, QC: Macdonald College) 47 (2): 14–42. Archived from the original on 26 May 2010. Retrieved 26 May 2010.
- Boase, C. (2001). "Bedbugs – back from the brink". Pesticide Outlook 12 (4): 159–162. doi:10.1039/b106301b. Retrieved 2010-05-27
- Szyndler, M.W.; Haynes, K.F.; Potter, M.F.; Corn, R.M.; Loudon, C. (2013). "Entrapment of bed bugs by leaf trichomes inspires microfabrication of biomimetic surfaces". Journal of The Royal Society Interface 10 (83). doi:10.1098/rsif.2013.0174. ISSN 1742-5662.
- Boase, Clive J. (April 2004). "Bed-bugs – reclaiming our cities". Biologist 51: 1–4. Retrieved 2010-06-07
- Potter, Michael F. (2011). "The History of Bed Bug Management – With Lessons from the Past". American Entomologist.
- Gerberg, Eugene J. (16 November 2008). "Entomologists in World War II". Proceedings of the DOD Symposium, ’Evolution of Military Medical Entomology’, Held 16 November 2008, Reno, NV. Annual Meeting of the Entomological Society of America (Armed Forces Pest Management Board). Retrieved 2013-11-11.
- Newsweek (8 September 2010). "The Politics of Bedbugs". Archived from the original on 21 October 2010. Retrieved 28 October 2010.
- Voiland, Adam (2007-07-16). "You May not be Alone". U.S. News & World Report 143 (2): 53–54.
- Megan Gibson (2010-08-19). "Are Bedbugs Taking Over New York City?". Time Magazine.
- Austin, James (2008). "Bed Bugs". Urban and Structural Pests. Center for Urban & Structural Entomology, Department of Entomology, Texas A&M. Retrieved 31 May 2010.
- Steelman, C.D.; Szalanski, A.L.; Trout, R.; McKern, J.A.; Solorzano, C.; Austin, J.W. (2008). "Susceptibility of the bed bug Cimex lectularius L. (Hemiptera: Cimicidae) to selected insecticides". Journal of Agricultural and Urban Entomology 25 (1): 41–51. doi:10.3954/1523-5475-25.1.41.
- Berg, Rebecca (2010). "Bed Bugs: The Pesticide Dilemma". Journal of Environmental Health 72 (10): 32–35. PMID 20556941.
- Oudhia P (1995). "Traditional knowledge about medicinal insects, mites and spiders in Chhattisgarh India". Insect Environment.
- Stephen L Doggett, Dominic E. Dwyer, Richard C Russell (January 2012). "Bed Bugs ClinicalRelevance and Control Options". Clinical Microbiology Review.
- "Online Etymology Dictionary". Etymonline.com. Retrieved 2013-11-11.
- "Bed Bug Home Page". Bedbug.org.au. 2005-10-14. Retrieved 2013-11-11.
- "Bed Bug Home Page". Bedbug.org.au. 2005-10-14. Retrieved 2013-11-11.
|Find more about Bed bugs at Wikipedia's sister projects|
|Definitions and translations from Wiktionary|
|Media from Commons|
|Learning resources from Wikiversity|
|Quotations from Wikiquote|
|Source texts from Wikisource|
|Textbooks from Wikibooks|
- bed bug on the University of Florida/IFAS Featured Creatures Web site
- Pollack, Richard; Alpert, Gary (2005). "Bedbugs: Biology and Management". Harvard School of Public Health. Archived from the original on 20 June 2010. Retrieved 2010-06-21.
- National Geographic segment on Bed bugs
- Bed Bug Fact Sheet highlights prevention tips as well as information on habits, habitat and health threats
- Bed bugs – University of Sydney and Westmead Hospital Department of Medical Entomology
- "Vector surveillance and control: Bed bug fact sheet" NYC Department of Health and Mental Hygiene 12 January 2008
- Understanding and Controlling Bed Bugs – National Pesticide Information Center
- CISR: Center for Invasive Species Research More information on Bed Bugs, with lots of photos and video