Ticks (Ixodida) are arachnids, typically 3 to 5 mm long, part of the superorder Parasitiformes. Along with mites, they constitute the subclass Acari. Ticks are external parasites, living by feeding on the blood of mammals, birds, and sometimes reptiles and amphibians. Ticks evolved by the Cretaceous period, the most common form of fossilisation being amber immersion. Ticks are widely distributed around the world, especially in warm, humid climates.

Temporal range: Cenomanian to present
Ixodes ricinus, a hard tick
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Subclass: Acari
Superorder: Parasitiformes
Order: Ixodida
Superfamily: Ixodoidea
Leach, 1815
  • Ixodidae – hard ticks
  • Argasidae – soft ticks
  • Nuttalliellidae – one species
18 genera, about 900 species

Almost all ticks belong to one of two major families, the Ixodidae or hard ticks, and the Argasidae or soft ticks. Adults have ovoid or pear-shaped bodies, which become engorged with blood when they feed, and eight legs. In addition to having a hard shield on their dorsal surfaces, hard ticks have a beak-like structure at the front containing the mouthparts, whereas soft ticks have their mouthparts on the underside of their bodies. Both families locate a potential host by odour or from changes in the environment.

Ticks have four stages to their lifecycle, namely egg, larva, nymph, and adult. Ixodid ticks have three hosts, taking at least a year to complete their lifecycle. Argasid ticks have up to seven nymphal stages (instars), each one requiring a blood meal. Because of their habit of ingesting blood, ticks are vectors of many diseases that affect humans and other animals.


Taxonomy and phylogeny

Fossilised tick in Dominican amber

Fossilized ticks are known from the Cretaceous onwards, most commonly in amber. They most likely originated in the Cretaceous (146 to 66 million years ago), with most of the evolution and dispersal occurring during the Tertiary (65 to 5 million years ago).[1] The oldest example is an argasid bird tick from Cretaceous New Jersey amber. The younger Baltic and Dominican ambers have also yielded examples that can be placed in living genera.[2] The tick Deinocroton draculi has been found with dinosaur feathers preserved in Cretaceous Burmese amber from 99 million years ago.[3]

Three families of ticks are described. The two large ones are the sister families of Ixodidae (hard ticks) and Argasidae (soft ticks). The third is the Nuttalliellidae, named for the bacteriologist George Nuttall. It comprises a single species, Nuttalliella namaqua,[4][5] and is the most basal lineage. It is found in southern Africa from Tanzania to Namibia and South Africa.[4][6] Ticks are closely related to the mites, within the subclass Acari.[7][8][9] rDNA analysis suggests that the Ixodidae are a clade, but that the Argasidae may be paraphyletic.[10][11]

The Ixodidae contain over 700 species of hard ticks with a scutum or hard shield, which the Argasidae lack. The Argasidae contain about 200 species; the genera accepted as of 2010 are Antricola, Argas, Nothoaspis, Ornithodoros, and Otobius.[4] They have no scutum, and the capitulum (mouth and feeding parts) is concealed beneath the body.[12]The phylogeny of the Ixodida within the Acari is shown in the cladogram, based on a 2014 maximum parsimony study of amino acid sequences of 12 mitochondrial proteins. The Argasidae appear monophyletic in this study.[13]

Ixodida (ticks)

Nuttalliellidae (1 species, southern Africa)

Ixodidae (hard ticks, ~700 species)

Argasidae (soft ticks, ~200 species)

Mesostigmata (mites, inc. Varroa)

Acariformes (mites)

Range and habitat

Tick species are widely distributed around the world,[14] but they tend to flourish more in countries with warm, humid climates, because they require a certain amount of moisture in the air to undergo metamorphosis, and because low temperatures inhibit their development from eggs to larvae.[15] Ticks are also widely distributed among host taxa, which include marsupial and placental mammals, birds, reptiles (snakes, iguanas, and lizards), and amphibians.[16] Ticks of domestic animals cause considerable harm to livestock by transmission of many species of pathogens, as well as causing anaemia and damaging wool and hides.[17] Some of the most debilitating species occur in tropical countries. Tropical bont ticks affect most domestic animals and occur in Africa and the Caribbean. The spinose ear tick has a worldwide distribution, the young feeding inside the ears of cattle and wild animals.[18]

In general, ticks are to be found wherever their host species occur. Migrating birds carry ticks with them on their journeys; a study of migratory birds passing through Egypt found more than half the bird species examined were carrying ticks. The species of tick often differed between the autumn and spring migrations, probably because of the seasonal periodicities of the different species.[19]

For an ecosystem to support ticks, it must satisfy two requirements; the population density of host species in the area must be high enough, and humidity must be high enough for ticks to remain hydrated.[20] Due to their role in transmitting Lyme disease, ixodid ticks, particularly the North American I. scapularis, have been studied using geographic information systems to develop predictive models for ideal tick habitats. According to these studies, certain features of a given microclimate – such as sandy soil, hardwood trees, rivers, and the presence of deer – were determined to be good predictors of dense tick populations.[21]

A habitat preferred by ticks is the interface where a lawn meets the woods,[22] or more generally, the ecotone, which is unmaintained transitional edge habitat between woodlands and open areas. Therefore, one tick management strategy is to remove leaf litter, brush, and weeds at the edge of the woods.[23] Ticks like shady, moist leaf litter with an overstory of trees or at least shrubs, and they deposit their eggs into such places in the spring, so that the larvae can emerge in the fall and crawl into low-lying vegetation. The 3 m of boundary closest to the lawn's edge are a tick migration zone, where 82% of tick nymphs in lawns are found.[24]

Anatomy and physiology

A hard-bodied tick of the family Ixodidae, the lone star tick

Ticks, like mites, are arthropods that have lost the segmentation of the abdomen that their ancestors had, with a subsequent fusion of the abdomen with the cephalothorax.[25] The tagmata typical of other Chelicerata have been replaced by two new body sections, the anterior capitulum (or gnathosoma), which is retractable and contains the mouthparts, and the posterior idiosoma, which contains the legs, digestive tract, and reproductive organs.[20] The capitulum is a feeding structure with mouthparts adapted for piercing skin and sucking blood; it is only the front of the head and contains neither the brain nor the eyes.[25] Features of the capitulum include the basis capitulum, two palps, two cutting chelicerae, and hypostome. The basis capitulum supports the rest of the feeding structures. Palps have a sensory role and are composed of three sections. The hypostome is used for blood extraction and is a hollow, tube-like structure.

The ventral side of the idiosoma bears sclerites, and the gonopore is located between the fourth pair of legs. In the absence of segmentation, the positioning of the eyes, limbs, and gonopore on the idiosoma provide the only locational guidance.[25]

Most ticks are inornate, meaning they are reddish or mahogany in color and lack markings. However, some species are ornate and have distinctive white patterns on the scutum. [26]

Larval ticks hatch with six legs, acquiring the other two after a blood meal and molting into the nymph stage.[27] In the nymphal and adult stages, ticks have eight legs, each of which has seven segments and is tipped with a pair of claws. The legs are sometimes ornamented and usually bear sensory or tactile hairs.[28] In addition to being used for locomotion, the tarsus of leg I contains a unique sensory structure, Haller's organ, which can detect odors and chemicals emanating from the host, as well as sensing changes in temperature and air currents.[29][30][31] Ticks can also use Haller's organs to perceive infrared light emanating from a host.[32] When not being used for walking, the legs remain tightly folded against the body.[29][30]

In nymphs and adults of the family Ixodidae, the capitulum is prominent and projects forwards from the body, a feature not present in the Argasidae. The eyes are close to the sides of the scutum, and the large spiracles are located just behind the coxae of the fourth pair of legs.[12] The hard, protective scutellum, characteristic of this family, covers the whole dorsal surface in males, but is restricted to a small, shield-like structure behind the capitulum in females and nymphs.[33] They differ, too, in their lifecycle; Ixodidae that attach to a host can bite painlessly and generally are unnoticed, and they remain in place until they engorge and are ready to change their skin; this process may take days or weeks. Some species drop off the host to moult in a safe place, whereas others remain on the same host and only drop off once they are ready to lay their eggs.[34]

A soft-bodied tick of the family Argasidae, beside eggs it has just laid

The body of the soft tick, family Argasidae, is pear-shaped or oval with a rounded anterior portion. The mouthparts cannot be seen from above, as they are on the ventral surface. The cuticle is leathery; often, a centrally positioned dorsal plate is seen, with ridges that project slightly above the surrounding surface, but with no decoration. A pattern of small, circular, depressed areas shows where muscles are attached to the interior of the integument. The eyes are on the sides of the body, the spiracles open between legs 3 and 4, and males and females only differ in the structure of the genital pore.[35]

The Nuttalliellidae can be distinguished from both ixodid and argasid ticks by a combination of a projecting capitulum at the front and a soft, leathery skin. Other distinguish characteristics include the position of the stigmata, the lack of setae, the strongly corrugated integument, and the form of the fenestrated plates.[36][37]

Ticks are extremely tough, hardy, and resilient animals. They can survive in a near-vacuum for as long as a half hour.[38] Their slow metabolism during their dormant periods enables them to go long periods between meals.[39] During droughts, they can endure dehydration without feeding for as long as 18 weeks, although ticks with limited energy reserves may succumb to desiccation after 36 weeks.[40] To keep from dehydrating, ticks hide in humid spots on the forest floor[41] or absorb water from subsaturated air by secreting hygroscopic fluid produced by the salivary glands onto the external mouthparts and then reingesting the water-enriched fluid.[42]

Ticks can withstand temperatures just above 0 °F (−18 °C) for more than two hours, and can survive temperatures in the 20–29 °F (−7 – −2 °C) range for at least two weeks. Ticks have even been found in Antarctica, where they feed on penguins.[43]

Diet and feeding

A questing tick, fingers for scale

Ticks satisfy all of their nutritional requirements as ectoparasites, feeding on a diet of blood. They are obligate hematophages, needing blood to survive and move from one stage of life to another. Ticks can fast for long periods, but eventually die if unable to find a host.[44] This behavior evolved approximately 120 million years ago through adaptation to blood-feeding.[45] The behavior evolved independently in the separate tick families, with differing host-tick interactions driving the evolutionary change.[45]

Some ticks attach quickly, while others wander around looking for thinner skin, such as is found on the ears of mammals. Depending on the species and life stage, preparing to feed can take from 10 minutes to two hours. On locating a suitable feeding spot, the tick grasps the host's skin and cuts into the surface.[44] They extract blood by cutting a hole in the host's epidermis, into which they insert their hypostome, and keep the blood from clotting by excreting an anticoagulant or platelet aggregation inhibitor.[46][47]

Ticks find their hosts by detecting animals' breath and body odors, or by sensing body heat, moisture, and vibrations.[48] They are incapable of flying or jumping, but many tick species, particularly Ixodidae, lie in wait in a position known as "questing". While questing, ticks cling to leaves and grasses by their third and fourth pairs of legs. They hold the first pair of legs outstretched, waiting to grasp and climb on to any passing host. Tick questing heights tend to be correlated with the size of the desired host; nymphs and small species tend to quest close to the ground, where they may encounter small mammalian or bird hosts; adults climb higher into the vegetation, where larger hosts may be encountered. Some species are hunters and lurk near places where hosts may rest. On receiving an olfactory or other stimulus, they crawl or run across the intervening surface.[48]

Other ticks, mainly the Argasidae, are "nidicolous", finding hosts in their nests or burrows, and in caves in the case of bats. They use the same stimuli as non-nidicolous species to identify hosts, with body heat and odors often being the main factors.[48] Many of them feed primarily on birds, though some Ornithodoros species, for example, feed on mammals. Both groups feed rapidly, typically biting painfully and drinking their fill within minutes. None of the species sticks to the host in the way that hard ticks do. Unlike the Ixodidae that have no fixed dwelling place except on the host, they live in sand or in crevices near animal dens or nests, or in human dwellings where they come out nightly to attack roosting birds, or emerge when they detect carbon dioxide in the breath of their hosts.[21]

In the Ixodidae, the tick stays in place until it is completely engorged. Its weight may increase by 200 to 600 times as compared to its weight before it started feeding. To accommodate this large expansion, cell division takes place and its cuticle grows larger; the tick may remain attached for days or weeks, depending on species, life stage, and host.[49] In the Argasidae, the tick's cuticle stretches to accommodate the fluid ingested, but does not grow new cells, with the weight of the tick increasing five- to 10-fold over the unfed state. The tick then drops off the host and typically remains in the nest or burrow until its host returns to provide its next meal.[35]

Tick saliva contains about 1,500 to 3,000 proteins depending on the tick species. The proteins with anti-inflammatory properties, called evasins, help ticks to feed for 8–10 days without being noticed by the host animal, as they block the host's chemokines and prevent painful inflammation. Researchers are studying these evasins with the goal of developing drugs to neutralise the chemokines that cause inflammation in myocarditis, heart attack, and stroke.[50]


Mites and nematodes feed on ticks, which are also a minor nutritional resource for birds. More importantly, they carry diseases as the primary hosts of pathogens such as spirochaetes, and without their agency, the organisms could not reach their secondary hosts. The diseases caused may debilitate their victims, and ticks may thus be assisting in controlling animal populations and preventing overgrazing.[51]

Certain infectious diseases of humans and other animals can be transmitted by ticks, with the species of tick involved tending to be those with a wide host range. Spread of disease in this way is enhanced by the extended time during which a tick remains attached, during which time the mobile host can be carried long distances, or in the case of bird hosts, across the sea. The infective agents can be present not only in the adult tick, but also in the eggs produced plentifully by the females. Many tick species have extended their ranges as a result of the movements of people, their pets, and livestock. With increasing participation in outdoor activities such as wilderness hikes, more people and their dogs may find themselves exposed to attack.[52]

Life cycle

Both ixodid and argasid ticks have four lifecycle stages: egg, larva, nymph, and adult.[53]


One-host ticks

Eggs laid in the environment hatch into larvae, which immediately seek out a host in which to attach and feed. Fed larvae molt into unfed nymphs that remain on the host. After engorging on the host's blood, the nymphs molt into sexually mature adults that remain on the host in order to feed and mate. Once a female is both fed and ready to lay eggs, only then does she drop off the host in search of a suitable area to deposit her eggs. Ticks that follow this life cycle are called one-host ticks. The winter tick Dermacentor albipictus and the cattle tick Boophilus microplus are examples of one-host ticks.[54]

Two-host ticks

Newly hatched larvae attach to a host in order to obtain a blood meal. They remain on the host after developing into nymphs. After emerging from their shed larval skins, the nymphs reattach to the host and feed. Once engorged, they drop off the host and find a safe area in the natural environment in which to molt into adults. Both male and female adults seek out a host on which to attach, which may be the same body that served as host during their early development. Once attached, they feed and mate. Females ready to lay eggs drop from the host to oviposit in the environment. Ticks that complete their life cycle in this manner are called two-host ticks, like Hyalomma anatolicum excavatum.[54]

Three-host ticks

Most ixodid ticks require three hosts, and their lifecycles take at least a year to complete. Thousands of eggs are laid on the ground by an adult female tick. When the larvae emerge, they attach and feed primarily on small mammals and birds. After feeding, they detach from their hosts and molt to nymphs on the ground, which then attach and feed on larger hosts before dropping off yet again in order to molt into adults. Adults seek out a third host on which to feed and mate. Female adults engorge on blood and prepare to drop off to lay her eggs on the ground, while males feed very little and remain on the host in order to continue mating with other females.[21][54]


Argasid ticks, unlike ixodid ticks, may go through up to seven nymphal stages (instars), requiring a meal of blood each time.[55] Their lifecycles range from months to years. The adult female argasid tick can lay a few hundred to over a thousand eggs over the course of her lifetime. Larvae feed very quickly and detach to molt into nymphs. Both male and female adults feed on blood, and they mate off the host. During feeding, any excess fluid is excreted by the coxal glands, a process that is unique to argasid ticks.[21]

Relationship with humans

Tick-borne disease

A sign in a Lithuanian forest warning of high risk of tick-borne encephalitis infection

Ticks are implicated in the transmission of a number of infections caused by pathogens such as bacteria, viruses, and protozoa. Sometimes, the tick harbours more than one type of pathogen, making diagnosis of the infection more difficult. Species of the bacterial genus Rickettsia are responsible for typhus, rickettsialpox, boutonneuse fever, African tick bite fever, Rocky Mountain spotted fever, Flinders Island spotted fever, and Queensland tick typhus (Australian tick typhus).[56] Other tick-borne diseases include Lyme disease and Q fever,[57] Colorado tick fever, Crimean Congo hemorrhagic fever, tularemia, tick-borne relapsing fever, babesiosis, ehrlichiosis, Bourbon virus, and tick-borne meningoencephalitis, as well as bovine anaplasmosis and probably the Heartland virus.[58]

Some species, notably the Australian paralysis tick, are also intrinsically venomous and can cause tick paralysis. Eggs can become infected with pathogens inside a female tick's ovaries, in which case the larval ticks are infectious immediately at hatching, before feeding on their first host.[55] Tropical bont ticks transmit the rickettsial disease heartwater, which can be particularly devastating in cattle.[18] The ticks carried by migratory birds may act as reservoirs and vectors of infectious diseases. Over 20 strains of pathogenic viruses were found in the autumn in the Egyptian migratory bird study.[19]

Not all ticks in an infective area are infected with pathogens, and both attachment of the tick and a long feeding session seem to be necessary for transmission of these diseases to take place.[52] Thus, tick bites often do not lead to infection, especially if the ticks are removed within 36 hours.[59] Adult ticks can be removed with fine-tipped tweezers or proprietary tick removal tools, disinfecting the wound.[60][61][62] It is also possible to freeze them off with a medical wart remover.[63] If the tick's head and mouthparts break off during removal, they can be removed with tweezers like a splinter. Ticks can be disposed of by flushing them down the toilet, placing them in a container of soapy water or alcohol, or sticking them to tape that can then be folded over and thrown away.[27][62]

Bifenthrin and permethrin, both pyrethroids, are sometimes used as tick-control measures, although a downside is that they are carcinogenic and can attack the nervous systems of other species besides ticks. Those who walk through tick-infested areas can make it harder for ticks to latch onto them by tucking their slacks into boots made of smooth rubber, which ticks have more trouble climbing than other material.[64][65]

Research since 2008 has documented red-meat allergies (known as Alpha-gal syndrome) in the U.S. due to lone star tick bites. The range of the problem has been expanding with the range of the tick. Other species of ticks are suspected of being responsible for meat allergies in other countries, including Sweden, Germany, and Australia.[66]

Many tick-transmitted viruses, such as Crimean-Congo hemorrhagic fever virus, Kyasanur Forest disease virus, Alkhumra hemorrhagic fever virus, and Omsk hemorrhagic fever virus, are classified as dangerous enough to require biosafety level 4 precautions in laboratory environments. This includes five levels of containment, viz., storage vials within humidified desiccators, within environmental chambers, within a tick suite, within a BSL4 laboratory. Precautions such as glove boxes, sticky pads, Vaseline barriers, safety suits, gloves, sticky tape, silicone vacuum grease, sticky trap paste, and micro mesh are used to safely contain ticks and prevent them from escaping.[67]

Population control measures

Researcher collecting ticks using the "tick dragging" method

With the possible exception of widespread DDT use in the Soviet Union, attempts to limit the population or distribution of disease-causing ticks have been quite unsuccessful.[68] The parasitoid chalcid wasp Ixodiphagus hookeri has been investigated for its potential to control tick populations. It lays its eggs into ticks;[69][lower-alpha 1] the hatching wasps kill their hosts.[70]

Predators and competitors of tick hosts can indirectly reduce the density of infected nymphs, thereby lowering tick-borne disease risk by lowering the density and/or tick burden of reservoir-competent hosts. A study in the Netherlands found that the number of larval ticks on bank voles and wood mice was lower at sites with significant red fox (Vulpes vulpes) and stone marten (Martes foina) activity.[71]

This supports the results of a study from the northeastern US, in which the incidence of Lyme borreliosis was negatively correlated with the density of red fox, which was suggested to be because foxes decrease the density of white-footed mice (Peromyscus leucopus), the most important reservoir-competent host for Borrelia burgdorferi in the US.[71][72]

Another natural form of control for ticks is the guineafowl, a bird species that consumes mass quantities of ticks.[73] Opossums also are net destroyers of ticks, killing around 90% of the ticks that attempt to attach and feed on them.[74] More generally, high animal diversity has a strongly protective effect against tick-borne disease.[24]

Topical tick medicines may be toxic to animals and humans. The synthetic pyrethroid insecticide phenothrin in combination with the hormone analogue methoprene was a popular topical flea and tick therapy for felines. Phenothrin kills adult ticks, while methoprene kills eggs. However, some products have been withdrawn, while others are known to cause adverse reactions.[75]

See also


  1. Micrographs of the wasp laying eggs into a tick, and the hole by which the young wasps emerge from the tick's dead body, are available in Plantard et al 2012.[69]


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