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''Cochliomyia macellaria'', also known as the secondary screwworm, is a species of blow fly in the family
Calliphoridae The Calliphoridae (commonly known as blow flies, blow-flies, carrion flies, bluebottles, greenbottles, or cluster flies) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing ba ...
. These screwworms are referred to as "secondary" because they typically infest wounds after invasion by primary
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 ...
-causing flies. While
blow flies The Calliphoridae (commonly known as blow flies, blow-flies, carrion flies, bluebottles, greenbottles, or cluster flies) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing b ...
may be found in every terrestrial
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 ...
, ''C. macellaria'' is primarily found in the United States, American
tropics The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referr ...
, and sometimes southern Canada. They are most common in the southeastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
in states like
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. ''C. macellaria'' have a
metallic Metallic may be a reference to: *Metal * Metalloid, metal-like substance *Metallic bonding, type of chemical bonding * Metallicity, in astronomy the proportion of elements other than helium and hydrogen in an object *Metallic color, a color that ...
greenish-blue
thorax The thorax or chest is a part of the anatomy of humans, 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 divisions of the cre ...
and a red-orange head and eyes. These adult blowflies range from 5–8 mm in size. Since the fly larvae infect the wounds and dead tissue of animals, these flies pose a grave medical and economic risk to humans and livestock. ''C. macellaria'' are attracted to carrion and garbage and are often found in slaughterhouses and outdoor markets in the tropics. While these flies carry many various types of '' Salmonella'' and viruses like the
swine influenza Swine influenza is an infection caused by any of several types of swine influenza viruses. Swine influenza virus (SIV) or swine-origin influenza virus (S-OIV) refers to any strain of the influenza family of viruses that is endemic in pigs. As ...
, ''C. macellaria'' can also serve as important
decomposer Decomposers are organisms that break down dead or decaying organisms; they carry out decomposition, a process possible by only certain kingdoms, such as fungi. Like herbivores and predators, decomposers are heterotrophic, meaning that they use o ...
s in our
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
.


Description

''Cochliomyia macellaria'' are classified as intermediate sized flies with a dull or bright metallic, blue-green coloration. On their
thorax The thorax or chest is a part of the anatomy of humans, 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 divisions of the cre ...
, there are three black
longitudinal Longitudinal is a geometric term of location which may refer to: * Longitude ** Line of longitude, also called a meridian * Longitudinal engine, an internal combustion engine in which the crankshaft is oriented along the long axis of the vehicl ...
stripes that interrupt the blue-green color. The eye and head of these flies are orange-red in color. ''C. macellaria'' also has pale
seta In biology, setae (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
e on the fronto-orbital plate outside of the frontal bristles. On ''C. macellaria'' the frontal row of bristles extend anteriorly to the base of the first antennal segment and consist of 12-14 bristles. The legs of the fly are orange brown to dark brown, and the coxae are orange brown to black with a green metallic luster. ''C. macellaria'' possess a dark reddish brown anterior
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
and an orange-red anterior
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
. The anterior tibia also has four short bristles that are placed on the
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
side. The color of the wings of the fly is orange brown tint towards the base. On these wings, the veins in the fly are orange brown to dark brown. The costal sessions 2 to 6 are in the proportions 78:56:96:30:6:6. The first genital section is black with a metallic green luster. On the fly, the scattered setae and a defined marginal row of bristles; the second segment is smaller with a dark brown to black tint with scattered setae.


Distinguishing features

''Cochliomyia macellaria'' is closely related to '' C. hominivorax'' but can be distinguished from ''C. hominivorax'' in three primarily ways: two of which are seen in adults and one which is seen in the larvae. In adults, the hairs on the fronto-orbital plate are black in C. ''hominivorax,'' but are pale in C. ''macellaria.'' Additionally, the central black stripe on the thorax extends only slightly in front of the mesonotal suture in ''C. hominivorax'' and well in front of the suture in ''C. macellaria''. ''C. macellaria'' larvae'','' unlike like ''C. hominivorax'' larvae'','' do not have pigmented tracheal trunks but instead have V-shaped spines on the
anal Anal may refer to: Related to the anus *Related to the anus of animals: ** Anal fin, in fish anatomy ** Anal vein, in insect anatomy ** Anal scale, in reptile anatomy *Related to the human anus: ** Anal sex, a type of sexual activity involving s ...
protuberance. ''C. macellaria'' also does not have an oral
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 ...
.


Distribution and territory

''Cochliomyia macellaria'' is most often found in the southeastern part of the United States, in states like
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
,
Louisiana Louisiana , group=pronunciation (French: ''La Louisiane'') is a state in the Deep South and South Central regions of the United States. It is the 20th-smallest by area and the 25th most populous of the 50 U.S. states. Louisiana is borde ...
, and
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
. Despite the high concentration in these states, these flies have been found as far north as Southern Canada and as south as the Neotropics, with the exclusion of countries like
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
and
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. While they have been found in this large expanse of areas, they have been shown to thrive most in warm and humid areas like the Southern United States, Caribbean Islands, Central America, and northern South America. In these areas, the population size typically increases during periods of extended rainfall. The territories of these blowflies are relatively small, especially during the time of
mating In biology, mating is the pairing of either opposite- sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reprod ...
. In this period, these flies will stay within a couple meters of one another.


Life cycle and history

The average life span of an adult ''C. macellaria'' is 2–6 weeks. In this time period, the females try to increase the chances of producing as many surviving offspring as possible. In a lifetime, ''C. macellaria'' may lay up to 1000 or more
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s. These eggs are typically laid in groups of 40 to 250. Females may also lay their eggs with other females, leading to an accumulation of thousands of eggs.


Eggs

The eggs of ''C. macellaria'' are laid in large groups (40–250 eggs at one time) and typically hatch in about 24 hours. The time taken for eggs to hatch often depends on moisture, temperature, and precipitation. In favorable conditions, the eggs may hatch in just four hours. These eggs are about 1 mm long and appear white to pearl white in color.


Larvae

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. ...
l stage of ''C. macellaria'' is referred to by the common name of secondary screwworms; this is due to the presence of small spines on each body segment that resemble parts of a screw. The larval stage of ''C. macellaria'' immediately follows the egg stage and is typically broken down into three substages or
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
s. Upon hatching, the larve appear a cream color, have cylindrical bodies with 10 or more spines around the spiracular area, possess incomplete peritremes (an indistinct or absent button), and have bands of small spines on each segment. ''C. macellaria,'' unlike like ''C. hominivorax,'' do not have pigmented tracheal trunks; instead, they have V-shaped spines on the anal protuberance. ''C. macellaria'' also does not have an oral sclerite. ''Cochliomyia macellaria'' larvae will feed on the decaying flesh of the animal that they have been laid on until they reach maturity. This stage of maturity is during the third instar and by this time point, the larvae may be as long as 17 mm. The entire larval stage is about four to seven days long, and afterwards, the larvae fall off the food source to pupate in the top layer of the soil.


Pupae

The ''C. macellaria'' larvae will typically burrow underneath the top layer of soil, leaves, garbage and begin to pupate there. During this stage, the outer layer begins to turn brown from the cream white that it used to be. The outer skin will begin to shrink and harden while the
pupa 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 thei ...
develops entirely in the hardened shell. Based on the temperature, the length of this stage will vary greatly. In warmer temperatures, the stage may last as short as seven days, but in colder
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measurement, measured with a thermometer. Thermometers are calibrated in various Conversion of units of temperature, temp ...
s, it may last as long as two months.


Adults

Adult ''C. macellaria'' are considered to be medium-sized
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced m ...
because they are about 6 to 9 mm in size and appear bright metallic blue-green to blueish purple in color. These adults typically spend one to two days maturing after leaving the pupae stage. Within four days of emerging from the
pupa 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 thei ...
stage, the adult ''C. macellaria'' become sexually mature and start looking for mates. The males will typically mate rapidly and will spend most of their time eating nearby
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 characte ...
and nectar from
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s. The females, on the other hand, will feed on the fluids of live wounds from animals. Contrary to the males, the females will travel long distances to find mates.


Food resources

Depending on the gender and stage in their development, ''C. macellaria'' will utilize different resources for energy and nutrients. During the larval stage, the ''C. macellaria'' will dig deeper into the
necrotic Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
wounds of their host animal and feed on the dead tissue. Both males and females will feast through this method. This time period is extremely crucial to the flies, as they must gather enough nutrients in order to last through the pupal stage without any food sources. This is one reason why the larvae are much larger than the size of the adult ''C. macellaria.'' As adults, the food specifications of the flies change. Female flies will continue to feed on tissues of animals; however, now they preferentially feed off of live tissue and tissue plasma. Males, at this point, will no longer consume tissue, but instead will eat nearby vegetation and intake nutrients from the nectar of flowers.


Mating

Although there are few studies on the mating patterns of ''C. macellaria,'' there is some information on the interactions between the males and females. Adult female ''C. macellaria'' have been observed to will release
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s that will stimulate the males
copulatory Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
attempts on contact. Even though depriving the ''C. macellaria'' adults of dietary
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s did not impact the potency of the female extracts, there was a reduced response in males for the pheromones.


Parental care

Male ''C. macellaria'' do not provide parental care. The males will typically mate quickly and have the females bear the
young Young may refer to: * Offspring, the product of reproduction of a new organism produced by one or more parents * Youth, the time of life when one is young, often meaning the time between childhood and adulthood Music * The Young, an American roc ...
. Since the females lay so many eggs, there is little care provided by the actual mothers after laying the eggs. The females do strategically lay the eggs in dead skin flesh so that once the eggs hatch into larvae, nutrition will be readily available.


Social behavior

Traditional social behavior like lekking has not been observed in ''C. macellaria;'' however, other social behaviors have been observed''.''


Egg laying

It is common for female ''C. macellaria'' to lay eggs with other females. Since females may lay up to 250 eggs, it is common for aggregations of thousands of eggs to infest entire animal carcasses.


Pheromone selectivity

While ''C. hominivorax'' is closely related to ''C. macellaria'', there are evolutionary developed methods that lead to reproductive isolation. One of these mechanisms is the interspecies response to the ''C. macellaria'' female
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
. Studies on newly colonized ''C. hoinivorax'' males have demonstrated that the males do not response to the ''C. macellaria'' pheromone.


Enemies


Larval predation

When studying the larval dispersal and predation for ''C. macellaria,'' studies have shown that there is interspecies competition and
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
. It has been demonstrated that '' C. albiceps'' larvae attack ''C. macellaria'' larvae during their dispersal process. Additionally, ''C. macellaria'' showed higher aggregation level in single than in double species. This may be explained by an
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary mechanism used to conserve and protect ''C. macellaria'' from predation.


Human depredation

Due to the medical and economic impact of the ''C. macellaria'' and other blowflies, an array of pesticides have been developed to reduce the ''C. macellaria'' population, including
pyrethrin The pyrethrins are a class of organic compounds normally derived from '' Chrysanthemum cinerariifolium'' that have potent insecticidal activity by targeting the nervous systems of insects. Pyrethrin naturally occurs in chrysanthemum flowers and i ...
aerosol.


Mimicry and protective coloration

While there are theories for protective coloration, there is no clear mimicry done by ''C. macellaria'' or by other insects specifically mimicking ''C. macellaria.'' The metallic green coloration may be a form of warding off predators, but this still in the process of being analyzed.


Impact


Medical

''Cochliomyia macellaria'' has extensive medical and economic implications. When ''C. macellaria'' larvae infest the dead and decaying tissues of animals or humans, myiasis may often occur. Once infestation occurs, a dark brown discharge will start to leaking from the wound. As the infestation increases, there is more agitation and inflamed tissue, which is accompanied by unpleasant smells. After the process of
clinical diagnosis Medical diagnosis (abbreviated Dx, Dx, or Ds) is the process of determining which disease or condition explains a person's symptoms and signs. It is most often referred to as diagnosis with the medical context being implicit. The information re ...
begins and the myiasis is recognized, then the larvae are easier to identify. Treatment of these infestations can be time intensive and leads to increased chances of reoccurrences. The first step is manual removal of the larvae followed by an antibiotic smear.


Economic

The
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animal ...
industry considers the secondary screwworm an important pest because of the enormous economic losses caused by cases of myiasis and disease transmission. The
medical Medicine is the science and practice of caring for a patient, managing the diagnosis, prognosis, prevention, treatment, palliation of their injury or disease, and promoting their health. Medicine encompasses a variety of health care practic ...
and pesticide treatment costs the United States millions of dollars annually.


Forensic importance

''Cochliomyia macellaria'' is the most common species of blow flies found on carrion in the southern United States. ''C. macellaria'' has recently gained recognition in forensic entomology because of its occurrence on decomposing remains. Since the analysis of the succession and occurrence has been well defined, it is possible to create postmortem interval estimations, which are crucial for forensic entomology. Adult ''C. macellaria'' in the southeastern United States are only attracted to the dead tissue on animals minutes after the death. In other regions of the United States, the adult flies are attracted to the dead flesh after a 24-hour delay.


References


Further reading

*


External links

* {{Taxonbar, from=Q13495225 Calliphoridae Articles created by Qbugbot Insects described in 1775 Taxa named by Johan Christian Fabricius Diptera of North America Insects of the United States