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''Cuterebra fontinella'', the mouse bot fly, is a species of New World skin bot fly in the family
Oestridae Botflies, also known as warble flies, heel flies, and gadflies, are a family of flies known as the Oestridae. Their larvae are internal parasites of mammals, some species growing in the host's flesh and others within the gut. '' Dermatobia hom ...
. ''C. fontinella'' is typically around in length with a black and yellow color pattern. ''C. fontinella'' develops by
parasitizing Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
nutrients from its host, typically the white-footed mouse. ''C. fontinella'' has even been known to parasitize humans in rare cases. Individuals parasitized by ''C. fontinella'' will develop a large bump on the skin that is indicative of parasitization.


Etymology

The genus name ''Cuterebra'' is a blend of the Latin words ''cutis'' : skin and ''terebra'' : borer with apparent shortening of expected ''Cutiterebra'' to ''Cuterebra''.


Distribution

''C. fontinella'' is found all around
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
, including in most of the continental United States, southern Canada, and northern Mexico. The prevalence of ''C. fontinella'' is dependant upon temperature. Colder regions will not be as densely populated as the warmer ones.


Habitat

''C. fontinella'' inhabits
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, ...
forests of North America. ''C. fontinella'' prefer territories near running water and with low-elevation vegetation. These flies are found in the highest density near the edges of their
habitats In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
.


Description

''C. fontinella'' look very similar to other species within the genus ''Cuterebra'' but have a few distinguishing features. The easiest indicator to use is the host species, as different species of Cuterebra infest different hosts. However, this method is not always reliable as ''C. fontinella'' have been known to infest multiple host species in addition to their preferred host, ''Peromyscus leucopus,'' the white-footed mouse. ''C. fontinella'' eggs are typically long and 0.03 mm wide. They are shaped like canoes and have a large groove along their underside; this groove enables attachment to vegetation. ''C. fontinella'' larvae transition from golden brown to black during development. The larvae are oval in shape. Grown larvae are typically long, wide, and thick. The larva is segmented into 12 sections, with backwards facing conical spines on almost every segment. The head segment is shield shaped, colored white or tan, contains antennal pits and retractable mandibles. The 12th segment bears two spiracles and is also lightly colored. However, the spines on this segment are outward facing. Adult ''C. fontinella'' have black hair on their
scutum The ''scutum'' (; plural ''scuta'') was a type of shield used among Italic peoples in antiquity, most notably by the army of ancient Rome starting about the fourth century BC. The Romans adopted it when they switched from the military formatio ...
and a black spot on their anipisterum. Adults are typically and have a close resemblance to
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
s.


Home range and territoriality

''C. fontinella'' flies are territorial insects. Males chase away intruding males while patrolling their territory by flying in figure-eight and oval patterns. Since females only fly when looking for a mate, the male tries to control as much high-quality territory as he can. They congregate above heat-reflecting surfaces on roadsides and near streams. These flies are very dependent on temperature; though they can live in a vast array of
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north ...
s, the
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
they live in influences their prevalence throughout the year.


Life history


Eggs

Egg development is slowed by temperatures below and under arid atmosphere conditions. The ideal conditions for development are warm and
humid Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity depe ...
, characteristic of southern climates. Stimulation from moisture and heat (often from the host passing by the egg) will cause the egg to hatch and the larva to be rubbed onto the host.


Larval instars

Once on the host, the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
enters through the nose, mouth, eyes, anus, or any open wound. Inside the body, the larva migrates and settles near the
groin In human anatomy, the groin (the adjective is ''inguinal'', as in inguinal canal) is the junctional area (also known as the inguinal region) between the abdomen and the thigh on either side of the pubic bone. This is also known as the medial comp ...
, then creates a “warble” with a small hole at the top layer of skin for breathing. The backward-facing spikes on the segments of the larva help to stabilize it.by preventing the larva from being pulled out of the host by gripping to the flesh surrounding it. The larvae typically emerge after the host dies (typically 3.5–4 weeks), and then burrow into the ground.


Pupae

''C. fontinella''
pupate A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
s in the ground and emerges as an adult. Diapause can occur during winter or other general unfavorable conditions. Pupae can remain in
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...
for as long as 12 months if need be. In simulated standard settings (27 °C and an alternating day and night light schedule), pupal development typically occurs in 50 days.


Adults

Adults of ''C. fontinella'' live a short life without feeding.
Oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...
development begins as soon as females leave the pupal stage of development. Oocyte development finishes in about 5 days. Mating can occur before oocyte development is finished. No secondary egg development occurs, likely due to the short adult lifespan of ''C. fontinella''. Female ''C. fontinella'' females typically lay their eggs in
vegetation Vegetation is an assemblage of plant species and the ground cover they provide. It is a general term, without specific reference to particular taxa, life forms, structure, spatial extent, or any other specific botanical or geographic charac ...
, especially that located near the homes/burrows of their intended hosts. Adult females can lay up to 2000 eggs.


Parasitism


Migration within the host

After entering the body, the larvae follow a distinct path within the host. If entering either through the nose, mouth, or eyes, the larvae first orient themselves using the host's nasal passageway. Then they travel through the upper
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gre ...
system, making their way to the
thoracic cavity The thoracic cavity (or chest cavity) is the chamber of the body of vertebrates that is protected by the thoracic wall (rib cage and associated skin, muscle, and fascia). The central compartment of the thoracic cavity is the mediastinum. There ...
. They then proceed to the
abdominal cavity The abdominal cavity is a large body cavity in humans and many other animals that contains many organs. It is a part of the abdominopelvic cavity. It is located below the thoracic cavity, and above the pelvic cavity. Its dome-shaped roof is th ...
and eventually to the
inguinal In human anatomy, the inguinal region refers to either the groin or the lower lateral regions of the abdomen. It may also refer to: * Conjoint tendon, previously known as the inguinal aponeurotic falx, a structure formed from the transversus abd ...
region of the host. If the larvae enter the host through an open wound or the anus, they find the closest portion of the standard tract, then continue their journey as they normally would.


Warbles

Once settled, the larva creates areas of swelling in the subcutaneous skin layer of their host. These swellings, known as warbles, are located between the anus and genital organs of the host. They last the same amount of time that the larva spends in its larval stage (3.5-4.0 weeks). The warble consists of a pore, a cavity, and a capsule. The pore serves as a breathing hole for the larva. As the larva grows, the size of the warble grows with it.
Translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable scattering of light. On a macroscopic scale (one in which the dimensions ...
yellow liquids are secreted from the larva. The cavity is the area of the warble where the larva lives. It expands gradually as the larva grows in size. The capsule is the tissue that surrounds the cavity. The capsule starts off as thin tissue, and thickens as larval development continues due to natural bodily healing of the host.


Effect on host


Effect on individual

Individuals infested with ''C. fontinella'' can experience
anemia Anemia or anaemia (British English) is a blood disorder in which the blood has a reduced ability to carry oxygen due to a lower than normal number of red blood cells, or a reduction in the amount of hemoglobin. When anemia comes on slowly, t ...
,
leukocytosis Leukocytosis is a condition in which the white cell (leukocyte count) is above the normal range in the blood. It is frequently a sign of an inflammatory response, most commonly the result of infection, but may also occur following certain parasi ...
, plasma protein imbalances, local tissue damage, and
splenomegaly Splenomegaly is an enlargement of the spleen. The spleen usually lies in the left upper quadrant (LUQ) of the human abdomen. Splenomegaly is one of the four cardinal signs of ''hypersplenism'' which include: some reduction in number of circulatin ...
. Infested female hosts produce fewer and smaller litters than those that are parasite-free. Counterintuitively, infestation actually leads to an increase in survival time for the host, not a decrease in survival as one would expect. This could be caused by the change in resource allocation from reproduction to body preservation, a change that is favorable for both the host's and the ''C. fontinella's'' survival. Interestingly, if multiple ''C. fontinella'' larvae infest a host simultaneously, the host instead experiences a decrease in survival time.


Host resistance

Resistance to infestation has been documented in hosts that have been previously infested. The resistance occurs at the entry points that have been used previously by larvae, as well as the genital region of the host where the larva typically creates the warble. Nasal, oral, and anal resistance cause a decrease in infestation rate when exposed to larvae of 15-30%. No resistance from repeated
ocular Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...
entry occurs. Maximum
antibody An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of t ...
production from the host occurs 28 days after infestation. The resistance causes the larvae to take abnormal alternate paths within the host. These abnormal paths caused the larvae to develop in atypical regions in the host. The
insecticide Insecticides are substances used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Insecticides are used in agriculture, medicine, industry and by consumers. Insecticides are claimed t ...
Ronnel can be used to effectively stop development of larvae by preventing the larvae from boring breathing holes in the skin of the host.


Effect on population

Infestation has varying effects on the host population's reproductive fitness, depending on the size of the habitat and the infestation rate of ''C. fontinella''. If the host population lives in a vast
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
with scattered areas of infestation, the reproductive rate of the host is essentially unaffected. However, in smaller habitats with higher and more uniform rates of infestation, reproductive rates can be noticeably decreased.


Food resources

The host serves as the primary food source. Different species of ''Cuterebra'' prey on various species of rodents. However, ''C. fontinella'' has been documented as preying on several different species of rodents. ''Peromyscus leucopus'' ( white-footed mouse) is the favored host for ''C. fontinella''. Typically, 19%-33% of all ''P. leucopus'' are infested within a year. Other hosts for ''C. fontinella'' include ''Lepus artemisia'' (
cottontail rabbit Cottontail rabbits are the leporid species in the genus ''Sylvilagus'', found in the Americas. Most ''Sylvilagus'' species have stub tails with white undersides that show when they retreat, giving them their characteristic name. However, this ...
), ''Ochrotomys nuttalli'' (
golden mouse The golden mouse (''Ochrotomys nuttalli'') is a species of New World mouse. It is usually 5–8 inches (12–25 cm) in body length, and has a soft pelage that ranges from golden-brownish to burnt orange in color. The genus name comes ...
), ''P. gossypinus'' (
cotton mouse The cotton mouse (''Peromyscus gossypinus'') is a species of rodent in the family Cricetidae found in the woodlands of the US South. Description Adults are about long, with a tail around , and weigh 34-51 g. Its general appearance is very sim ...
), ''P. maniculatus'' (
eastern deer mouse ''Peromyscus maniculatus'' is a rodent native to eastern North America. It is most commonly called the eastern deer mouse; when formerly grouped with the western deer mouse (''P. sonoriensis''), it was referred to as the North American deerm ...
), ''Heteromys irroratus'' (
mexican spiny pocket mouse The Mexican spiny pocket mouse (''Heteromys irroratus'') is a species of rodent in the family Heteromyidae. It is native to Mexico and Texas in the United States where it is found in dry, scrubby habitats. The IUCN has assessed it as being of "le ...
), and even very rarely humans. The highest infestation rates occur in late summer and early fall.


Mating

Adults aggregate in open, sunny areas to find their mates. However, ''C. fontinella'' does not mate at temperatures below 20 °C. Adult males fly 1–2 m above the ground for up to 4 hours a day in the presence of sunlight to attract a mate. Once a female demonstrates her interest, the pair finds a nearby branch or leaf for stability. The female grabs the branch or leaf and the male mounts her. The pair copulates for about 3 minutes. Ovipositor The
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
of ''C. fontinella'' is unusually short relative those of the other members of the family Oestridae. Covered by dense hairs, the ovipositor resembles a
horseshoe A horseshoe is a fabricated product designed to protect a horse hoof from wear. Shoes are attached on the palmar surface (ground side) of the hooves, usually nailed through the insensitive hoof wall that is anatomically akin to the human ...
, with two
sclerite A sclerite (Greek , ', meaning " hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly ...
ends and a
chitinous Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chiti ...
plate between them.


Social behavior

Males of ''C. fontinella'' fly around their territories from the time at which the temperature exceeds 20 °C to the early afternoon. Flight is initiated only during sunlight, but can continue during brief cloud coverage not exceeding 15 minutes. Since their flight is limited by temperature, the amount of time spent flying is dependent on the weather and temperature of the day. Females of ''C. fontinella'' are only seen flying when searching for a mate. Males are extremely territorial, and usually occupy a stretch of territory 17 m long, along the bank of a stream. They chase most airborne intruders that travel through their territory, regardless of their species. If two males intersect in the air, they grab onto each other and tumble to the ground. Chases persist for 10–15 minutes. ''C. fontinella'' tolerates a population density of up to 250 flies/km2. Fly density varies depending on weather conditions and the time of the year.


Genetic identification

Genetic analysis can also be used to discriminate between species. COI and COII genes are reliable markers to differentiate between species. Using species specific markers, scientists can accurately identify the species of a botfly at any stage of life. In many cases larvae look ambiguous enough to be confused for another species, so genetic identification is very important. Hybridization between species within the genus ''Cuterebra'' has been known to occur, and can cause ambiguity within testing.


Subspecies

The two subspecies of ''C. fontinella'' are ''C. f. fontinella'' and ''C. f. grisea'' (deer mouse bot fly).


Interactions with humans

Rare cases of ''C. fontinella'' host infestations have been reported, but are not the norm. In most cases, the larvae remain in relatively benign locations such as in the eye or in the subcutaneous regions within the eyelid. Occasionally, however, the larvae gain access to the tracheal-pulmonary system. Consequent symptoms in the human host include cold-like symptoms and flares of coughing. The larvae are expelled from the human host when the host coughs out a bloody secretion containing the larva. Another species of bot fly called ''
Dermatobia hominis The human botfly, ''Dermatobia hominis'' (Greek δέρμα, skin + βίος, life, and Latin '' hominis'', of a human), is a species of botfly whose larvae parasitise humans (in addition to a wide range of other animals, including other primates ...
'' (human botfly) commonly infests humans in Central and South America. Most cases of human infestation within North America are caused by the victim traveling into regions where ''D. hominis'' is present. As of 1989, 55 cases of
myiasis Myiasis is the parasitic infestation of the body of a live animal by fly larvae (maggots) which grow inside the host while feeding on its tissue. Although flies are most commonly attracted to open wounds and urine- or feces-soaked fur, some spe ...
caused by species within the ''Cuterebra'' genus have been documented. Treatment typically consists of removal of the larva and then prevention of secondary bacterial infections. If the warble is accessible, one can remove the larva by depositing petroleum jelly over the breathing hole of the parasite; this causes the larva to emerge for air and enable easier removal. Larvae within or near the eye sometimes require surgery for removal. Larvae that die within the
vitreous humor The vitreous body (''vitreous'' meaning "glass-like"; , ) is the clear gel that fills the space between the lens and the retina of the eyeball (the vitreous chamber) in humans and other vertebrates. It is often referred to as the vitreous humor ...
of the eye do not need to be removed, they are broken down and absorbed by natural chemical processes within the host.


Conservation

A major threat to infestation rates of ''C. fontinella'' is pasture burning. When soil reaches high temperatures, pupating larvae die, and ash produced during burning causes
microclimate A microclimate (or micro-climate) is a local set of atmospheric conditions that differ from those in the surrounding areas, often with a slight difference but sometimes with a substantial one. The term may refer to areas as small as a few squ ...
changes. These changes contribute significantly to increasing greater congregation of ''C. fontinella''.


References


Further reading

* * *


External links

* {{Taxonbar, from=Q13505862 Oestridae Articles created by Qbugbot Insects described in 1827