Chrysomya Putoria
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''Chrysomya putoria'', also known as the tropical African latrine blowfly, is a fly species belonging to the blowfly family,
Calliphoridae The Calliphoridae (commonly known as blowflies, blow flies, blow-flies, carrion flies, bluebottles, or greenbottles) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing bait, ...
''.'' ''C. putoria'' is native to
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
and has recently spread to the Americas. These flies pose significant health risks, especially due to their close association with human settlements. Adult flies can carry pathogens, while larvae may cause
myiasis Myiasis ( ), also known as flystrike or fly strike, is the parasitic infestation of the body of a live animal by fly larvae (maggots) that grow inside the host while feeding on its Biological tissue, tissue. Although flies are most commonly attr ...
by growing and feeding on the flesh of domestic animals and humans. Other myiasis-causing flies in the same genus are '' C. bezziana'' and '' C. megacephala''. ''C. putoria'' and other flies that feed on decomposing tissue are used as important tools in
forensic entomology Forensic entomology is a branch of applied entomology that uses insects and other arthropods as a basis for legal evidence. Insects may be found on cadavers or elsewhere around crime scenes in the interest of forensic science. Forensic entom ...
to establish the
post-mortem interval The post-mortem interval (PMI) is the time that has elapsed since an individual's death. When the time of death is not known, the interval may be estimated, and so an approximate time of death established. Postmortem interval estimations can ra ...
, or the time elapsed since death.


Description

The adult body is long, slightly larger than the average
housefly The housefly (''Musca domestica'') is a fly of the suborder Cyclorrhapha. It possibly originated in the Middle East, and Cosmopolitan distribution, spread around the world as a commensal of humans. Adults are gray to black, with four dark, lo ...
, and metallic green, with an occasional tinge of metallic dark blue or bronze. The face of ''C. putoria'' is generally dark in color. Their wings are translucent, and the posterior edge of the abdomen is striped with black bands. While both sexes have black
frons Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insec ...
, the female has a much broader frons located between its widely-spaced eyes. This species has easily visible dusting on the upper side of its
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
, allowing it to be distinguished from many other closely related Calliphoridae species.


Habitat and distribution

''C. putoria'' originates from the tropics of the
Old World The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
, specifically Africa. The fly is widely distributed across the central and southern regions of the African continent, and found in countries including Zambia, Ethiopia, and Madagascar. Within the past few decades, several ''Chrysomya'' species, including ''C. putoria,'' invaded South America. These flies may have been initially introduced into Brazil from Africa in the 1970s before spreading across the rest of the continent. This dispersal may be connected to human mobility through the use of transportation like airplanes and ships. ''C. putoria'' is now found in many South American countries, including Argentina, Brazil, Colombia, Paraguay, and Peru. This spread of wild fly species may have also been facilitated by the fly's strong flying skills and synanthropy, known as the ability to benefit by closely associating with human habitats. As a tropical species, ''C. putoria'' prefers to inhabit warmer climates. In its native Old World habitat, this fly is generally found in areas with a temperature above 26.7 °C.


Taxonomy

Members of the family Calliphoridae lay
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e in the tissue and
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
of
warm-blooded Warm-blooded is a term referring to animal species whose bodies maintain a temperature higher than that of their environment. In particular, homeothermic species (including birds and mammals) maintain a stable body temperature by regulating ...
animals, such as in
pit latrine A pit latrine, also known as pit toilet, is a type of toilet that collects human waste in a hole in the ground. Urine and feces enter the pit through a drop hole in the floor, which might be connected to a toilet seat or squatting pan for user ...
s. The African blowfly ''C. chloropyga'' is the
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''C. putoria'' and also breeds on feces and decaying flesh. Due to morphological similarities, these flies were previously considered to be subspecies.
Phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis of
cytochrome c oxidase The enzyme cytochrome c oxidase or Complex IV (was , now reclassified as a translocasEC 7.1.1.9 is a large transmembrane protein complex found in bacteria, archaea, and the mitochondria of eukaryotes. It is the last enzyme in the Cellular respir ...
gene sequences indicates that the two species diverged just a few thousand years ago. ''C. albiceps and C. putoria'' both spread to the
Peruvian Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long and wide (widest between 18°S ...
by 1980. While they broadly overlap at the elevation of , their distributions indicate differential temperature preferences. ''C. putoria'' prefers inhabiting warmer, tropical areas below , while ''C. albiceps'' can typically be found in temperate areas between .


Life history


Egg

''C. putoria'' eggs are approximately long, rod-shaped, and cream-white in color.


Larva

''C. putoria'' undergoes three larval instar stages. After hatching from the egg, the larva in the first instar is long with 11 spine bands aligned along its 12 separate body segments. In addition, a pair of spiracles can be seen on its most posterior segment with a singular spiracular opening. During the second instar, the larva remains cream-white in color and grows to in length. It develops fleshy folds and a pair of spiracles along its 2nd segment. The posterior spiracles now present two spiracular openings. The maxillary palp and antennae are also more developed. In the final instar stage, the larva reaches a length of and develops larger fleshy folds. The posterior spiracles are then fully developed with three spiracular openings. The spine on the anal segment has a bell-shaped appearance.


Pupa

The
pupa A pupa (; : 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 life cycle, the stages th ...
l stage lasts approximately 5 days. Early on, the malleable outer layer of the final instar becomes brittle and hard. The surface of the pupa is rough, bumpy, and unreflective. At the same time, the translucent cuticle gains creamy-white pigmentation. The body transforms on the second day, taking on yellow pigmentation and increasing the size of the head and wings. On the third day, the eyes turn orange and the body pigmentation shows a reddish tinge with a black stripe across the abdomen. On the fourth day, three black stripes appear across the abdomen, the wings, the legs become black, and the eyes and body darken into a brown color. On the fifth and final day, the body turns black, and the fly ecloses by breaking through the pupal casing.


Adult

The adult female has approximately 80
ovariole An ovariole is a tubular component of the insect ovary, and the basic unit of egg production. Each ovariole is composed of a germarium (the germline Stem-cell niche, stem cell niche) at the anterior tip, a set of developing Oocyte, oocytes containe ...
s in each of her two
ovaries The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are endocr ...
. She also has a pair of tubular sexual accessory glands that lead into the common
oviduct The oviduct in vertebrates is the passageway from an ovary. In human females, this is more usually known as the fallopian tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, or will dege ...
. In addition, the female has three spherical
spermatheca The spermatheca (pronounced : spermathecae ), also called ''receptaculum seminis'' (: ''receptacula seminis''), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, Oligochaeta worms and certain other in ...
e that store sperm from males after mating.


Food resources

Adult ''C. putoria'' females lay eggs on feces, decaying flesh, and rotting food matter. As a result, the larvae feed and grow on these substrates. If there is a scarcity of resources, ''C. putoria'' larvae may
cannibalize Cannibalism is the act of consuming another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded in more than 1,500 species. Human cannibalism is also well document ...
one another after 24 hours. In addition, injured larvae leak
hemolymph Hemolymph, or haemolymph, is a fluid, similar to the blood in invertebrates, that circulates in the inside of the arthropod's body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph c ...
, which may induce cannibalistic tendencies in other larvae.


Social behavior

Similar to other blowfly species, ''C. putoria'' exhibit larval gregariousness. The aggregation of larvae in isolated, temporal patches of food demonstrates
intraspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
and
interspecific competition Interspecific competition, in ecology, is a form of competition in which individuals of ''different'' species compete for the same resources in an ecosystem (e.g. food or living space). This can be contrasted with mutualism, a type of symbiosis. ...
.


Predators and pathogens

''Podisus nigrispinus'', a species of stinkbug, is a predator of ''C. putoria'' larvae. The larvae of ''C. albiceps'', another member of the blowfly family'','' are facultative predators of ''C. putoria'' larvae and the larvae of other fly species. ''C. albiceps'' larvae will surround then fatally pierce the target. ''C. putoria'' are affected by pathogenic fungi ''
Metarhizium anisopliae ''Metarhizium anisopliae'' is a fungus, the type species in the ''Metarhizium'' genus. It grows naturally in soils throughout the world and causes disease in various insects by acting as a parasitoid. Ilya Ilyich Mechnikov, Ilya I. Mechnikov nam ...
'' and ''
Beauveria bassiana ''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the group of entomopathogenic fungi. It is used as a ...
'' that act as
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s and can eventually kill.


Genetics

The
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
chromosome number is 2n = 12, and all the chromosomal pairs are metacentric. Their count consists of 5 autosomal pairs and a sex chromosomal pair with an
XY sex-determination system The XY sex-determination system is a sex-determination system present in many mammals (including humans), some insects (''Drosophila''), some snakes, some fish (guppy, guppies), and some plants (''Ginkgo'' tree). In this system, the sex of an ...
.
Karyotype A karyotype is the general appearance of the complete set of chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is discerned by de ...
analysis indicates that the pericentromeric region of the autosomal chromosomes includes a C band, while the 3rd pair of chromosomes additionally includes an interstitial band.


Physiology

Studies on the effect of temperature changes on the development of these flies found that the optimal temperature for egg survivability is between , consistent with the flies' preference for tropical climates. ''C. putoria'' has a developmental threshold of ~, which is relatively higher than ''C. chloropyga's'' threshold of ~, supporting their differential adaptations to temperature.


Interactions with humans


Medical significance

The synanthropy of these flies is especially concerning, as their preference for human settlements may contribute to a major health impact in developing countries. Since these flies often breed in latrines, they are especially dangerous pests in areas with low sanitation. The effect of these flies is further compounded by their high population density and capability of widespread dispersal. Breeding in the feces of domestic animals, especially poultry, can spread pathogens to humans. Furthermore, their attraction to breeding on human and dog feces can make them vectors of diarrheal pathogens. This is supported by PCR analysis showing ''C. putoria'' flies infected with ''
E. coli ''Escherichia coli'' ( )Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. is a gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus ''Escherichia'' that is commonly foun ...
'', ''
Salmonella ''Salmonella'' is a genus of bacillus (shape), rod-shaped, (bacillus) Gram-negative bacteria of the family Enterobacteriaceae. The two known species of ''Salmonella'' are ''Salmonella enterica'' and ''Salmonella bongori''. ''S. enterica'' ...
'', and ''
Shigella ''Shigella'' is a genus of bacteria that is Gram negative, facultatively anaerobic, non–spore-forming, nonmotile, rod shaped, and is genetically nested within ''Escherichia''. The genus is named after Kiyoshi Shiga, who discovered it in 1 ...
''.
Myiasis Myiasis ( ), also known as flystrike or fly strike, is the parasitic infestation of the body of a live animal by fly larvae (maggots) that grow inside the host while feeding on its Biological tissue, tissue. Although flies are most commonly attr ...
can also occur if an adult female lays eggs inside the body of a living animal. While chemical
insecticide Insecticides are pesticides used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. The major use of insecticides is in agriculture, but they are also used in home and garden settings, i ...
s are commonly used to control synanthropic fly populations, there are potential adverse side-effects of these chemicals on the health of humans, animals, crops, and the environment. Efforts to target latrines may be effective at reducing the impact of these pests, since latrines are a major source of fly populations. ''Chrysomya putoria'' larvae have potential use in
maggot therapy Maggot debridement therapy (also known as MDT, larval therapy, or simply maggot therapy) is a type of biotherapy involving the introduction of live, disinfected maggots (fly larvae) into non-healing skin and soft-tissue wounds of a human or othe ...
since their removal of dead tissue in wounds can promote the growth of new tissue.


Forensic importance

Since the larvae of ''C. putoria'' and other blowfly species actively decompose
cadaver A cadaver, often known as a corpse, is a Death, dead human body. Cadavers are used by medical students, physicians and other scientists to study anatomy, identify disease sites, determine causes of death, and provide tissue (biology), tissue to ...
s, understanding their life cycle duration can be used to estimate the PMI (post-mortem interval). Cocaine and other drugs in the cadaver's system can accelerate the life cycle of the larvae, therefore introducing error into PMI analysis. Ongoing studies on the impact of drugs on the life cycle duration can reduce potential sources of PMI estimation errors.


References

{{Taxonbar, from=Q3675990 Calliphoridae Insects described in 1830 Parasitic flies Taxa named by Christian Rudolph Wilhelm Wiedemann Diptera of Africa