Glossina fuscipes
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''Glossina fuscipes'' is a riverine fly species in the genus ''
Glossina Tsetse ( , or ) (sometimes spelled tzetze; also known as tik-tik flies), are large, biting flies that inhabit much of tropical Africa. Tsetse flies include all the species in the genus ''Glossina'', which are placed in their own family, Glos ...
'', which are commonly known as
tsetse flies Tsetse ( , or ) (sometimes spelled tzetze; also known as tik-tik flies), are large, biting flies that inhabit much of tropical Africa. Tsetse flies include all the species in the genus ''Glossina'', which are placed in their own family, Glos ...
. Typically found in sub-Saharan Africa but with a small Arabian range, ''G. fuscipes'' is a regional
vector Vector most often refers to: *Euclidean vector, a quantity with a magnitude and a direction *Vector (epidemiology), an agent that carries and transmits an infectious pathogen into another living organism Vector may also refer to: Mathematic ...
of African trypanosomiasis, commonly known as sleeping sickness, that causes significant rates of morbidity and mortality among
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s and
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 ...
. Consequently, the species is among several being targeted by researchers for
population control Population control is the practice of artificially maintaining the size of any population. It simply refers to the act of limiting the size of an animal population so that it remains manageable, as opposed to the act of protecting a species from ...
as a method for controlling the disease.


Physical Description

''G. fuscipes'' are often brown or grey-brown in color. Their bodies tend to have varied dark and light patches, effectively
camouflaging Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
them on surfaces such as bark, rock, or soil. At rest, ''G. fuscipes'' appear slim as they fold their wings on their backs so that one lays on top of the other. This is in contrast to
houseflies The housefly (''Musca domestica'') is a fly of the suborder Cyclorrhapha. It is believed to have evolved in the Cenozoic Era, possibly in the Middle East, and has spread all over the world as a commensal of humans. It is the most common f ...
and
blowflies 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 ...
whose wings project outward at an angle while resting on their backs. Following a blood meal, the insect's abdomen will appear large, rounded and red.


Males

When the male ''G. fuscipes'' is examined from the
ventral side Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
, a rounded structure named the hypopygium can be seen at the posterior end of the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. Immediately in front of the hypopygium is a plate bearing dark hairs called hectors. Both the hypopygium and the hectors help distinguish male from females and serve to grasp onto the end of the female abdomen during
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 ...
. As copulation commences, the hypopygium unfolds to uncover superior and inferior
clasper In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's c ...
s as well as the aedeagus.


Females

The end of the female abdomen lacks any significant structures that would be the counterpart of the male hypopygium and hectors; however, females display a
vulva The vulva (plural: vulvas or vulvae; derived from Latin for wrapper or covering) consists of the external female sex organs. The vulva includes the mons pubis (or mons veneris), labia majora, labia minora, clitoris, vestibular bulbs, vulv ...
, which can exhibit several small plates that aid in species identification.


Life Cycle


Egg stage

In a few hours, the sperm move from the spermatophore into the spermatheca, where they remain active for the remainder of the female's life. Eggs are fertilized immediately as they enter the uterus by sperm from the spermatheca that come into contact and penetrate the anterior portion of the egg. The fertilized egg remains in the uterus for about four days as the instar larva begins to develop. Once she has mated, a female can produce larvae for the remainder of her life. At about 25 °C, a female fly will produce mature larva every 9–10 days with the exception of the first, which may take up to 18–20 days. Lower temperatures result in a lower rate of breeding whereas higher temperatures increase the rate of breeding. Temperatures either too high or too low may cease breeding altogether.


Larva

The ''G. fuscipes''
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. ...
in passes through three
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 as it grows up to when the fully grown larva is dropped by a female fly. The larva has a mouth at the anterior end and two spiracles at the posterior end. Rather unusually, the larva spends most of its time and does all its feeding within the mother's body. Apart from food stored in the egg, the food supply for the three larval instar stages comes from the mother's milk gland. The milky secretions of this gland are expelled out of the gland duct at the head end of the larva. The larva sucks up the milky secretion and passes it directly to the
midgut The midgut is the portion of the embryo from which most of the intestines develop. After it bends around the superior mesenteric artery, it is called the "midgut loop". It comprises the portion of the alimentary canal from the end of the foregut a ...
where it is slowly digested and assimilated. For air supply, the larva depends on air entering the vulva of the female. The air must pass into the female's posterior spiracles or polypneustic lobes to reach the larva.


Abortion

If a larva fails to reach its full size, it will be prematurely expelled from the uterus. The aborted larva dies. Abortions could be due to the mother fly not obtaining enough food or also when carelessly handled or exposed to insecticide. Eggs are subjected to abortion for the same reasons.


Pupa

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 ...
is a dark brown, shorter than the larva that produced it, and rounded with polypneustic lobes at the posterior end. The lobes are distinctively shaped and can help to distinguish the ''G. fuscipes'' pupa from that of other flies. The pupa also has a hard case on its outside called the puparium. The pupal stage lasts about four to five weeks according to temperature. Higher temperatures shorten the pupal period. In contrast, lower temperatures lengthen the pupal period to more than 50 days in certain climates. However, temperature extremes will cause death.


Adult

When ready to emerge, the young adult fly expands its
ptilinum The ptilinum is an eversible pouch on the head, above the base of the antenna in schizophoran flies (a section Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, ...
to burst open puparium's end. Out of the fresh hole and surrounding soil, the adult emerges by using the ptilinum, struggling to the top of the soil and into open air. At this stage, the adult's body is very soft while its wings are small and crumpled. After a few urinations, the wings will expand towards their proper size. From the time between the emergence of the fly and its first meal, the adult is called a teneral fly. After the first blood meal has been taken, the fly is then termed a non-teneral fly.


Reproduction


Mating

During mating, males settle on the back of the female. Claspers at the posterior end of the male abdomen unfold in order to grip the end of the female abdomen. This mating position may be maintained for an hour or two before the duo parts. Females typically mate a young age, either before or around the same time of their first blood meals. Females usually mate only once in their lives though it is possible mate more than once, whereas males tend to mate several times. Older males are more likely to mate successfully than very young males. During mating, the aedeagus is inserted into the
vulva The vulva (plural: vulvas or vulvae; derived from Latin for wrapper or covering) consists of the external female sex organs. The vulva includes the mons pubis (or mons veneris), labia majora, labia minora, clitoris, vestibular bulbs, vulv ...
and reaches into the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The ...
as far as the
spermatheca The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other in ...
exit. A sizable ball of sperm is deposited there in the form of a
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
. At the conclusion of mating, the male releases his grip on the female before flying away.


Distribution and Habitat

''G. fuscipes'' are found in sub-Saharan African and a subpopulation of '' G. f. fuscipes'' exists in the very southern part of the Arabian peninsula. ''G. f. f.'' and '' G. m. submorsitans'' are the only subspecies of ''Glossina'' which survive outside Africa, including in southwestern
Saudi Arabia Saudi Arabia, officially the Kingdom of Saudi Arabia (KSA), is a country in Western Asia. It covers the bulk of the Arabian Peninsula, and has a land area of about , making it the fifth-largest country in Asia, the second-largest in the A ...
. They prefer high-humidity areas, namely biomes such as mangrove swamps, rain forests, lake shores, and gallery forests along rivers. ''G. fuscipes'' occupies a large inland block centered on
Democratic Republic of the Congo The Democratic Republic of the Congo (french: République démocratique du Congo (RDC), colloquially "La RDC" ), informally Congo-Kinshasa, DR Congo, the DRC, the DROC, or the Congo, and formerly and also colloquially Zaire, is a country in ...
but also covering land surrounding countries in addition to
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the nort ...
,
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the C ...
and a southern portion of Chad.


Evolution and taxonomy

The genus
Glossina Tsetse ( , or ) (sometimes spelled tzetze; also known as tik-tik flies), are large, biting flies that inhabit much of tropical Africa. Tsetse flies include all the species in the genus ''Glossina'', which are placed in their own family, Glos ...
is regarded as an isolated genus and it is usually classified into its own family Glossinidae. The genus is further divided into three subgenera, Morsitans, Fusca, and Palpalis, the latter of which being the subgenus to which ''G. fuscipes'' belongs. The species is further broken down into subspecies '' G. f. fuscipes'', '' G. f. martinii'', and '' G. f. quanzensis''.


Food Resources

''G. fuscipes'' feed on vertebrate blood and have been traditionally described as strictly
hematophagous Hematophagy (sometimes spelled haematophagy or hematophagia) is the practice by certain animals of feeding on blood (from the Greek words αἷμα ' "blood" and φαγεῖν ' "to eat"). Since blood is a fluid tissue rich in nutritious p ...
. Glucose sugars are not a metabolic requirement for this species because it uses a proline-alanine shuttle system for the distribution of energy. Instead, triglycerides are used for storage in fly body fat and milk secretions. However, researchers have conducted laboratory experiments and a field study that show ''G. fuscipes'' are able to feed on sugar water in the lab and wild flies contain sugar residues. Although continuous feeding with high sugar concentrations appeared to be
toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subs ...
, sugar given either occasionally or at low concentrations did not affect mortality and
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
.


Predators

''G. fuscipes'' adults and pupae are a food source for a variety of
predators 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 ...
including
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
s and
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s. However, no insectivorous species is known to solely feed on ''G. fuscipes'' or tsetse flies in general. Thus, a reduction in insectivorous birds during general tsetse fly control campaigns could be attributed to the simultaneous insecticide-related removal of other insect species than decreases in tsetse flies themselves.


Trypanosomiasis

Some trypanosome species, transmitted by ''G. fuscipes'' and other tsetse fly species, cause the
infectious disease An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable di ...
trypanosomiasis Trypanosomiasis or trypanosomosis is the name of several diseases in vertebrates caused by parasitic protozoan trypanosomes of the genus ''Trypanosoma''. In humans this includes African trypanosomiasis and Chagas disease. A number of other diseas ...
. In humans, ''G. fuscipes'' trypanosomiasis is also known as
sleeping sickness African trypanosomiasis, also known as African sleeping sickness or simply sleeping sickness, is an insect-borne parasitic infection of humans and other animals. It is caused by the species ''Trypanosoma brucei''. Humans are infected by two typ ...
. In animals, the disease may be known as ''
nagana Animal trypanosomiasis, also known as nagana and nagana pest, or sleeping sickness, is a disease of vertebrates. The disease is caused by trypanosomes of several species in the genus ''Trypanosoma'' such as ''Trypanosoma brucei''. '' Trypanosom ...
'' or '' surra'' according to the animal species infected as well as the trypanosome species involved. ''Nagana'' typically refers to the disease specifically in cattle and horses; however, it is commonly used to describe any type of animal trypanosomiasis.


Disease vectors and hosts

''G. fuscipes'', alongside other tsetse flies, are prominent biological vectors of protozoan
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
belonging to the genus ''
Trypanosoma ''Trypanosoma'' is a genus of kinetoplastids (class Trypanosomatidae), a monophyletic group of unicellular parasitic flagellate protozoa. Trypanosoma is part of the phylum Sarcomastigophora. The name is derived from the Greek ''trypano-'' ( ...
'' known to cause the namesake diseases in various vertebrate species including
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s, antelopes, bovine cattle, camels,
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million yea ...
s,
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to domesticated ...
,
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s, and
pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
s. The parasites are transmitted to humans via bites from ''G. fuscipes'', which have acquired their infection from other human beings or animals harboring human-pathogenic parasites. The table below summarizes this information for the ''G. fuscipes'' species; however, the diseases listed below may be transferred by other
tsetse fly Tsetse ( , or ) (sometimes spelled tzetze; also known as tik-tik flies), are large, biting flies that inhabit much of tropical Africa. Tsetse flies include all the species in the genus ''Glossina'', which are placed in their own family, Glos ...
species in addition to ''G. fuscipes''.


Population control

The containment of
sleeping sickness African trypanosomiasis, also known as African sleeping sickness or simply sleeping sickness, is an insect-borne parasitic infection of humans and other animals. It is caused by the species ''Trypanosoma brucei''. Humans are infected by two typ ...
and
nagana Animal trypanosomiasis, also known as nagana and nagana pest, or sleeping sickness, is a disease of vertebrates. The disease is caused by trypanosomes of several species in the genus ''Trypanosoma'' such as ''Trypanosoma brucei''. '' Trypanosom ...
would be of great benefit to rural communities in sub-Saharan Africa, alleviating poverty and improving food security, thus efforts are undertaken in rein in local populations of ''G. fuscipes'' via methods such as pesticide campaigns,
trapping Animal trapping, or simply trapping or gin, is the use of a device to remotely catch an animal. Animals may be trapped for a variety of purposes, including food, the fur trade, hunting, pest control, and wildlife management. History Neolithi ...
, or the
sterile insect technique The sterile insect technique (SIT) is a method of biological insect control, whereby overwhelming numbers of sterile insects are released into the wild. The released insects are preferably male, as this is more cost-effective and the females ma ...
.


Mutualism


Microbiome

''G. fuscipes'' flies rely on the obligate
symbiont Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
l genus '' Wigglesworthia'' to supplement their diets with nutrients essential for fecundity. The adult
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splint ...
relies similarly on ''Wigglesworthia'' for activation and development. A secondary, facultative symbiont is the genus ''
Sodalis ''Sodalis'' is a genus of bacteria within the family Pectobacteriaceae. This genus contains several insect endosymbionts and also a free-living group. It is studied due to its potential use in the biological control of the tsetse fly. ''Sodalis' ...
'', which is present in tsetse populations considered to play a role in the ability to transmit trypanosomes. Finally, the third symbiont is the genus ''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reproducti ...
'', transovarially transmitted between generations. To enhance transmission and survival, ''Wolbachia'' has
evolved 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 ...
mechanisms to alter host reproduction. Using both culture-dependent and independent methods, it was shown that Kenyan populations of the subspecies ''G. f. fuscipes'' harbor diverse range of bacteria. Of the flies tested, bacteria were isolated from 72% of the sample population with 23 bacterial species identified. Of these, the
Bacillota The Bacillota (synonym Firmicutes) are a phylum of bacteria, most of which have gram-positive cell wall structure. The renaming of phyla such as Firmicutes in 2021 remains controversial among microbiologists, many of whom continue to use the earl ...
phylum constituted 16 species, seven of which belong to the genus ''
Bacillus ''Bacillus'' (Latin "stick") is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum '' Bacillota'', with 266 named species. The term is also used to describe the shape (rod) of other so-shaped bacteria; and the plural ''Bacill ...
''.


See also

*
List of diseases spread by invertebrates Invertebrates are very common vectors of disease. A vector is an organism which spreads disease from one host to another. Invertebrates spread bacterial, viral and protozoan pathogens by two main mechanisms. Either via their bite, as in the case of ...


References


External links

* {{Taxonbar, from=Q14601440 Diptera of Africa Insect vectors of pathogens Hippoboscoidea