Flea, the common name for theorderSiphonaptera, includes 2,500 species of small flightlessinsects that live as externalparasites ofmammals andbirds. Fleas live byingesting the blood of their hosts. Adult fleas grow to about 3 millimetres (1⁄8 inch) long, are usually dark in color, and have bodies that are "flattened" sideways or narrow, enabling them to move through their hosts' fur or feathers. They lack wings; their hind legs are extremely well adapted for jumping. Their claws keep them from being dislodged, and their mouthparts are adapted for piercing skin and suckingblood. Some species can leap 50 times their body length, a feat second only to jumps made by another group of insects, thesuperfamily offroghoppers. Flea larvae are worm-like, with no limbs; they have chewing mouthparts and feed on organic debris left on their hosts' skin.
Genetic evidence indicates that fleas are a specialised lineage of parasiticscorpionflies (Mecoptera)sensu lato, most closely related to the familyNannochoristidae. The earliest known fleas lived in the MiddleJurassic; modern-looking forms appeared in theCenozoic. Fleas probably originated on mammals first and expanded their reach to birds. Each species of flea specializes, more or less, on one species of host: many species of flea never breed on any other host; some are less selective. Some families of fleas are exclusive to a single host group; for example, the Malacopsyllidae are found only onarmadillos, theIschnopsyllidae only onbats, and the Chimaeropsyllidae only onelephant shrews.
Fleas are wingless insects,1.5 to 3.3 millimetres (1⁄16 to1⁄8 inch) long, that are agile, usually dark colored (for example, the reddish-brown of thecat flea), with aproboscis, or stylet, adapted to feeding by piercing the skin and sucking their host's blood through their epipharynx. Flea legs end in strong claws that are adapted to grasp a host.[1]
Unlike other insects, fleas do not possesscompound eyes but instead only have simple eyespots with a single biconvex lens; some species lack eyes altogether.[2] Their bodies are laterally compressed, permitting easy movement through the hairs or feathers on the host's body. The flea body is covered with hard plates called sclerites.[1] These sclerites are covered with many hairs and short spines directed backward, which also assist its movements on the host. The tough body is able to withstand great pressure, likely anadaptation to survive attempts to eliminate them by scratching.[3]
Fleas lay tiny, white, oval eggs. The larvae are small and pale, have bristles covering their worm-like bodies, lack eyes, and have mouth parts adapted to chewing. The larvae feed on organic matter, especially the feces of mature fleas, which contain dried blood. Adults feed only on fresh blood.[4]
Their legs are long, the hind pair well adapted for jumping; a flea can jump vertically up to 18 centimetres (7 inches) and horizontally up to 33 cm (13 in),[5] making the flea one of the best jumpers of all known animals (relative to body size), second only to thefroghopper. A flea can jump 60 times its length in height and 110 times its length in distance, equivalent to a 1.8 m (6 ft) adult human jumping 110 m (361 ft) vertically and 200 m (656 ft) horizontally. Rarely do fleas jump from dog to dog. Most flea infestations come from newly developed fleas from the pet's environment.[6] The flea jump is so rapid and forceful that it exceeds the capabilities of muscle, and instead of relying on direct muscle power, fleas store muscle energy in a pad of the elastic protein namedresilin before releasing it rapidly (like a human using a bow and arrow).[7] Immediately before the jump, muscles contract and deform the resilin pad, slowly storing energy which can then be released extremely rapidly to power leg extension for propulsion.[8] To prevent premature release of energy or motions of the leg, the flea employs a "catch mechanism".[8] Early in the jump, the tendon of the primary jumping muscle passes slightly behind the coxa-trochanter joint, generating atorque which holds the joint closed with the leg close to the body.[8] To trigger jumping, another muscle pulls the tendon forward until it passes the joint axis, generating the opposite torque to extend the leg and power the jump by release of stored energy.[8] The actual take off has been shown by high-speed video to be from the tibiae and tarsi rather than from thetrochantera (knees).[7]
Fleas areholometabolous insects, going through the fourlifecycle stages ofegg,larva,pupa, andimago (adult). In most species, neither female nor male fleas are fully mature when they first emerge but must feed on blood before they become capable of reproduction.[3] The first blood meal triggers the maturation of the ovaries in females and the dissolution of the testicular plug in males, and copulation soon follows.[9] Some species breed all year round while others synchronise their activities with their hosts' life cycles or with local environmental factors and climatic conditions.[10] Flea populations consist of roughly 50% eggs, 35% larvae, 10% pupae, and 5% adults.[5]
The number of eggs laid depends on species, with batch sizes ranging from two to several dozen. Thetotal number of eggs produced in a female's lifetime (fecundity) varies from around one hundred to several thousand. In some species, the flea lives in the host's nest or burrow and the eggs are deposited on the substrate,[9] but in others, the eggs are laid on the host itself and can easily fall off onto the ground. Because of this, areas where the host rests and sleeps become one of the primaryhabitats of eggs and developing larvae. The eggs take around two days to two weeks to hatch.[5]
Flea larvae emerge from the eggs to feed on any available organic material such as dead insects, faeces,conspecific eggs, and vegetable matter, some species are subdermal endoparasites on their parent’s hosts. In laboratory studies, some dietary diversity seems necessary for proper larval development. Blood-only diets allow only 12% of larvae to mature, whereas blood and yeast or dog chow diets allow almost all larvae to mature.[11] Another study also showed that 90% of larvae matured into adults when the diet included nonviable eggs.[12] They are blind and avoid sunlight, keeping to dark, humid places such as sand or soil, cracks and crevices, under carpets and in bedding.[13] The entire larval stage lasts between four and 18 days.[14]
Given an adequate supply of food, larvaepupate and weavesilken cocoons after three larval stages. Within the cocoon, the larva molts for a final time and undergoes metamorphosis into the adult form. This can take just four days, but may take much longer under adverse conditions, and there follows a variable-length stage during which the pre-emergent adult awaits a suitable opportunity to emerge. Trigger factors for emergence include vibrations (including sound), heat (in warm-blooded hosts), and increased levels ofcarbon dioxide, all of which may indicate the presence of a suitable host.[5] Large numbers of pre-emergent fleas may be present in otherwise flea-free environments, and the introduction of a suitable host may trigger a mass emergence.[13]
Once the flea reaches adulthood, its primary goal is to find blood and then to reproduce.[15] Female fleas can lay 5000 or more eggs over their life, permitting rapid increase in numbers.[16] Generally speaking, an adult flea only lives for 2 or 3 months. Without a host to provide a blood meal, a flea's life can be as short as a few days. Under ideal conditions of temperature, food supply, and humidity, adult fleas can live for up to a year and a half.[16] Completely developed adult fleas can live for several months without eating, so long as they do not emerge from theirpuparia. Optimum temperatures for the flea's life cycle are 21 to 30 °C (70 to 86 °F) and optimum humidity is 70%.[17]
Adult female rabbit fleas,Spilopsyllus cuniculi, can detect the changing levels ofcortisol andcorticosterone hormones in the rabbit's blood that indicate it is getting close to giving birth. This triggers sexual maturity in the fleas and they start producing eggs. As soon as the baby rabbits are born, the fleas make their way down to them and once on board they start feeding, mating, and laying eggs. After 12 days, the adult fleas make their way back to the mother. They complete this mini-migration every time she gives birth.[17]
Between 1735 and 1758, the Swedish naturalistCarl Linnaeus first classified insects, doing so on the basis of their wing structure. One of the seven orders into which he divided them was "Aptera", meaning wingless, a group in which as well as fleas, he includedspiders,woodlice andmyriapods. It wasn't until 1810 that the French zoologistPierre André Latreille reclassified the insects on the basis of their mouthparts as well as their wings, splitting Aptera intoThysanura (silverfish),Anoplura (sucking lice) and Siphonaptera (fleas), at the same time separating off the arachnids andcrustaceans into their own subphyla.[18] The group's name, Siphonaptera, is zoological Latin from the Greeksiphon (a tube) andaptera (wingless).[19]
It was historically unclear whether the Siphonaptera are sister to theMecoptera (scorpionflies and allies), or are inside that clade, making "Mecoptera" paraphyletic. The earlier suggestion that the Siphonaptera are sister to theBoreidae (snow scorpionflies)[20][21][22] is not supported. A 2020 genetic study recovered Siphonaptera within Mecoptera, with strong support, as the sister group toNannochoristidae, a small,relictual group of mecopterans native to the Southern Hemisphere. Fleas and nannochoristids share several similarities with each other that are not shared with other mecopterans, including similar mouthparts as well as a similar sperm pump organisation.[23]
Relationships of Siphonaptera per Tihelka et al. 2020.[23]
Fleas likely descended fromscorpionflies, insects that are predators or scavengers.[23] Fossils of large, winglessstem-group fleas with siphonate (sucking) mouthparts from the MiddleJurassic[24] to EarlyCretaceous have been found in northeastern China and Russia, belonging to the familiesSaurophthiridae andPseudopulicidae, as well asTarwinia from the Early Cretaceous of Australia. Most flea families formed after the end of the Cretaceous (in thePaleogene and onwards). Modern fleas probably arose in the southern continental area ofGondwana, and migrated rapidly northwards from there. They most likely evolved withmammal hosts, only later moving tobirds.[25]
Siphonaptera is a relatively small order of insects: members of the order undergo complete metamorphosis and are secondarily wingless (their ancestors had wings which modern forms have lost). In 2005, Medvedev listed 2005 species in 242 genera, and despite subsequent descriptions of new species, bringing the total up to around 2500 species,[20] this is the most complete database available. The order is divided into four infraorders and eighteen families. Some families are exclusive to a single host group; these include the Malacopsyllidae (armadillos), Ischnopsyllidae (bats) and Chimaeropsyllidae (elephant shrews).[26]
Many of the known species are little studied. Some 600 species (a quarter of the total) are known from single records. Over 94% of species are associated withmammalian hosts, and only about 3% of species can be considered to be specific parasites ofbirds. The fleas on birds are thought to have originated from mammalian fleas; at least sixteen separate groups of fleas switched to avian hosts during the evolutionary history of the Siphonaptera. Occurrences of fleas on reptiles is accidental, and fleas have been known to feed on thehemolymph (bloodlike body fluid) ofticks.[26]
Flea phylogeny was long neglected, the discovery ofhomologies with the parts of other insects being made difficult by their extreme specialization. Whiting and colleagues prepared a detailed molecular phylogeny in 2008, with the basic structure shown in the cladogram. TheHectopsyllidae, including the harmfulchigoe flea or jigger, is sister to the rest of the Siphonaptera.[20]
As of 2023[update], there are 21 recognized families within the order Siphonaptera, 3 of which are extinct.[27] In addition, some researchers have suggested that the subfamilyStenoponiinae should be elevated to its own family (Stenoponiidae).[28]
Fleas feed on a wide variety ofwarm-bloodedvertebrates including dogs, cats, rabbits, squirrels, ferrets, rats, mice, birds, and sometimes humans. Fleas normally specialise in one host species or group of species, but can often feed but not reproduce on other species.Ceratophyllus gallinae affects poultry as well as wild birds.[29] As well as the degree of relatedness of a potential host to the flea's original host, it has been shown that avian fleas that exploit a range of hosts, only parasitise species with low immune responses. In general, host specificity decreases as the size of the host species decreases. Another factor is the opportunities available to the flea to change host species; this is smaller in colonially nesting birds, where the flea may never encounter another species, than it is in solitary nesting birds. A large, long-lived host provides a stable environment that favours host-specific parasites.[30]
Although there are species named dog fleas (Ctenocephalides canis Curtis, 1826) and cat fleas (Ctenocephalides felis), fleas are not always strictly species-specific. A study inVirginia examined 244 fleas from 29 dogs: all were cat fleas. Dog fleas had not been found in Virginia in more than 70 years, and may not even occur in the US, so a flea found on a dog in the United States is likely a cat flea (Ctenocephalides felis).[31][32]
One theory of human hairlessness is that the loss of hair helped humans to reduce their burden of fleas and other ectoparasites.[33]
In many species, fleas are principally a nuisance to their hosts, causing anitching sensation which in turn causes the host to try to remove the pest by biting, pecking or scratching. Fleas are not simply a source of annoyance, however. Flea bites cause a slightly raised, swollen, irritating nodule to form on the epidermis at the site of each bite, with a single puncture point at the centre, like amosquito bite.[34]: 126 This can lead to aneczematous itchy skin disease calledflea allergy dermatitis, which is common in many host species, including dogs and cats.[29] The bites often appear in clusters or lines of two bites, and can remain itchy and inflamed for up to several weeks afterwards. Fleas can lead to secondary hair loss as a result of frequent scratching and biting by the animal. They can also causeanemia in extreme cases.[34]: 126
John Donne's erotic metaphysical poem "The Flea", published in 1633 after his death, uses theconceit of a flea, which has sucked blood from the male speaker and his female lover, as an extendedmetaphor for their sexual relationship. The speaker tries to convince a lady to sleep with him, arguing that if the mingling of their blood in the flea is innocent, then sex would be also.[42]
The comic poemSiphonaptera, written in 1915 by the mathematicianAugustus De Morgan, describes an infinite chain of parasitism made of ever larger and ever smaller fleas.[43]
Flea circuses provided entertainment to nineteenth century audiences. These circuses, extremely popular in Europe from 1830 onwards, featured fleas dressed as humans or towing miniature carts,chariots,rollers orcannon. These devices were originally made bywatchmakers orjewellers to show off their skill at miniaturization. A ringmaster called a "professor" accompanied their performance with a rapid circus patter.[44][45]
Oriental rat fleas,Xenopsylla cheopis, can carry thecoccobacillusYersinia pestis. The infected fleas feed on rodent vectors of this bacterium, such as theblack rat,Rattus rattus, and then infect human populations with theplague, as has happened repeatedly from ancient times, as in thePlague of Justinian in 541–542.[46] Outbreaks killed up to 200 million people across Europe between 1346 and 1671.[47] TheBlack Death pandemic between 1346 and 1353 likely killed over a third of the population of Europe.[48]
The bankerCharles Rothschild devoted much of his time toentomology, creating a large collection of fleas now in the Rothschild Collection at theNatural History Museum, London. He discovered and named theplaguevector flea,Xenopsylla cheopis, also known as the oriental rat flea, in 1903.[50] Using what was probably the world's most complete collection of fleas of about 260,000 specimens (representing some 73% of the 2,587 species and subspecies so far described), he described around 500 species and subspecies of Siphonaptera. He was followed in this interest by his daughterMiriam Rothschild, who helped to catalogue his enormous collection of the insects in seven volumes.[51][52]
Fleas have a significant economic impact. In America alone, approximately $2.8 billion is spent annually on flea-related veterinary bills and another $1.6 billion annually for flea treatment with pet groomers. Four billion dollars is spent annually for prescription flea treatment and $348 million for flea pest control.[13]
^abcHinkle, Nancy C.; Koehler, Philip G. (2008). "Cat Flea, Ctenocephalides felis felis Bouché (Siphonaptera: Pulicidae)". In Capinera, John L. (ed.).Encyclopedia of Entomology. Springer Netherlands. pp. 797–801.doi:10.1007/978-1-4020-6359-6_536.ISBN978-1-4020-6242-1.
^Meyer, John R. (28 March 2016)."Siphonaptera". North Carolina State University. Archived fromthe original on 25 September 2022. Retrieved3 December 2016.
^Wiegmann, Brian; Yeates, David K. (2012).The Evolutionary Biology of Flies. Columbia University Press. p. 5.ISBN978-0-231-50170-5.Recently, a close affinity between Siphonaptera and Mecoptera has been convincingly demonstrated via morphology (Bilinski et al. 1998) and molecular data (Whiting 2002), rendering Mecoptera paraphyletic, but making the clade including Mecoptera and Siphonaptera monophyletic
^Zurita, A.; Callejón, R.; De Rojas, M.; Gómez López, M.S.; Cutillas, C. (December 2015). "Molecular study of Stenoponia tripectinata tripectinata (Siphonaptera: Ctenophthalmidae: Stenoponiinae) from the Canary Islands: taxonomy and phylogeny".Bulletin of Entomological Research.105 (6):704–711.doi:10.1017/s0007485315000656.PMID26282009.S2CID35756267.
^Neri, Janice (2008). "Between Observation and Image: Representations of Insects in Robert Hooke's Micrographia". In O'Malley, Therese; Meyers, Amy R. W. (eds.).The Art of Natural History. National Gallery of Art. pp. 83–107.ISBN978-0-300-16024-6.
^Roncalli, Amici R. (June 2004). "La storia della pulce nell'arte e nella letteratura" [The history of the flea in art and literature].Parasitologia (in Italian).46 (1):15–18.PMID15305680. See also the2009 version.
^Black, Joseph, ed. (2010).The Broadview Anthology of British Literature, Volume 2 (2nd ed.). Broadview Press.ISBN978-1-55481-290-5.
^Schmäschke, R. (1 April 2000). "The flea in cultural history and first effects of its control".Berliner und Münchener Tierärztliche Wochenschrift.113 (4):152–160.ISSN0005-9366.PMID10816916.
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