Lutzomyia

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Lutzomyia
Temporal range: Burdigalian - Recent
Lutzomyia longipalpis-sandfly.jpg
Lutzomyia longipalpis taking a blood meal
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Psychodidae
Subfamily: Phlebotominae
Genus: Lutzomyia
França, 1924
Species

Lutzomyia is a genus of phlebotomine sand flies consisting of nearly 400 species, [1] at least 33 of which have medical importance as vectors of human disease. [2] Species of the genus Lutzomyia are found only in the New World, distributed in southern areas of the Nearctic and throughout the Neotropical realm. [2] Lutzomyia is one of the two genera of the subfamily Phlebotominae to transmit the Leishmania parasite, with the other being Phlebotomus , found only in the Old World. Lutzomyia sand flies also serve as vectors for the bacterial Carrion's disease and a number of arboviruses. [1]

Contents

Evolution

The genus, named after Adolfo Lutz, is known from the extinct Burdigalian (20–15  mya) species Lutzomyia adiketis found as a fossil in Dominican amber on the island of Hispaniola. [3] It is thought that species in the genus Lutzomyia all originated in the lowland forests to the east of the Andes mountain range, and that their radiation throughout the Neotropics was sparked by dry periods of the Pleistocene, driving colonisation further north and west to areas of higher humidity and leading to reproductive isolation. [4]

Systematics

The classification of species within the genus Lutzomyia is largely unresolved, and relies on often controversial divisions based on morphological taxonomic characters. Such analyses can suffer from polymorphisms within a species, the existence of cryptic species and the frequent lack of distinct morphological characters amongst females. Research has begun in an attempt to resolve evolutionary relationships between species in the genus, using molecular methods to create phylogenies based on ribosomal DNA sequences. [2]

Biology

The Lutzomyia sand flies are small, biting insects with a body length of up to only 3 mm. They are hairy insects with their colour ranging from a near-white to near-black. [5]

Life stages

Life stageCharacteristic features [1]
EggsFemale Lutzomyia sand flies will typically lay small batches of around 40-70 dark, elliptical eggs within a moist microhabitat. For most species, this will require the female to take a bloodmeal prior to oviposition. However, there are some species of the genus which are able to lay the first batch of eggs without a bloodmeal.
LarvaeLarval development of the Lutzomyia sand fly involves at least 18 days of development through four instars, before reaching maturity. Following hatching, the first instar larva will feed on any readily available organic matter. Development through the four instars produces a small, mature larva which is caterpillar-like in appearance. This larva will then aim to find a slightly drier environment in which to pupate.
PupaeThe pupal stage of development typically involves between 7 and 12 days of resting prior to emergence of the adults. It is generally the case that adult males emerge first.
AdultsEmergence of adult Lutzomyia sand flies occurs and, in males, is followed by sexual maturation within 24 hours. This sexual maturation is marked by the 180° rotation of the male's external genitalia. Adult males will then seek to mate, generally by using pheromones to find a resting site with potential mates. Males exhibit courtship behaviour, such as by varying wing beat patterns.

Feeding

Only the female Lutzomyia sand flies are blood-feeding, requiring the bloodmeal to provide the nutrients for the maturation of eggs. Therefore, it is only the females who have medical importance as vectors of disease. Both males and females require carbohydrate food as adults, the source of which remains unclear. [1]

The most medically important species for human disease transmission are those which predominantly favour humans as a source of bloodmeal. These species are known as anthropophillic and tend to feed at around dusk. However, there are examples of anthropophillic species that will attack in the daytime. The majority of anthropohillic Lutzomyia species in the Americas are exophillic, which means that they favour biting outside of homes. [1]

Resting

The resting behaviour of Lutzomyia sand flies, like many other aspects of their biology, is important to understand for targeted, vector-based control methods to reduce transmission of Leishmaniasis. For example, residual spraying of insecticides can be targeted at known resting sites to increase effectiveness. Lutzomyia sand flies generally rest outdoors, with the type of resting site varying between species and in response to the seasons and the availability of particular microhabitats. The largest resting microhabitat is the forest floor, but sand flies will also rest in a variety of other areas, such as in the nests and burrows of mammals, within the trunks of hollow trees and inside bat caves. [1] The resting position adopted by Lutzomyia sand flies is characteristic to the genus, with wings angling above the abdomen. [5]

Medical importance

Leishmaniasis

The sand fly genus Lutzomyia includes all species responsible for transmission, in the New World, of the Leishmania parasite, the causative agent of leishmaniasis. Of the more than 350 Lutzomyia species identified, less than 10% are known or suspected to transmit leishmaniasis to humans. [6] The disease is in endemic in 22 countries of tropical and subtropical America, where it is generally considered a zoonosis. [7] That is, the parasite cycles between mammalian reservoir hosts and the sand flies, with humans serving as incidental, dead-end hosts. Common reservoirs in the Americas include a number of rodent species, as well as dogs, sloths and armadillos. [7] The Lutzomyia sand flies maintain transmission amongst reservoir species and allow the disease to 'jump' to humans, causing the visceral, cutaneous or mucocutaneous forms of the disease depending on the Leishmania species. Visceral Leishmaniasis is a serious form of the disease in particular that affects the internal organs. This specific disease is primarily transmitted by the species Lutzomyia longipalpis . [8] The human-sand fly-human cycle of transmission, known as anthroponotic, is limited to two Leishmania species endemic in the Old World and so does not involve Lutzomyia sand flies. [7]

Giemsa-stained amastigotes of Leishmania (Viannia) panamensis Giemsa-Stained Amastigotes of Leishmania (Viannia) panamensis.jpeg
Giemsa-stained amastigotes of Leishmania (Viannia) panamensis

The New World Leishmania species transmitted by Lutzomyia sand flies are often classified in the subgenus, Viannia. [9] These genetically distinct parasites show markedly different patterns of development within the New World sand flies when compared to those seen in the Old World Phlebotomus sand flies. The first stages of development are similar between all Leishmania species, with the sandfly taking up the amastigote form of the parasite following a bite of an infected host. However, unlike in the Old World, the replicated parasites then migrate to the hindgut of the Lutzomyia sand fly, a feature which is thought to be essential in allowing the parasite to become established. [10] The parasite then undergoes further development into the infective, promastigote stage as it migrates to anterior end of the insect, ready for inoculation into a new, susceptible host.[ citation needed ]

The high level of species diversity in the genus Lutzomyia, coupled with its high adaptability is leading to increasing risk of Leishmania transmission in the New World in response to global change. Lutzomyia sand flies transmit the disease in the tropics and subtropics, regions which are subject to high levels of deforestation associated with continual development. Deforestation, with the establishment of settlements at the periphery of primary or secondary forest, increases the risk of Leishmania transmission by creating a selection pressure for the adaptation of sand flies to these new peridomestic environments. [11] In southern Brazil, for example, levels of cutaneous leishmaniasis are increasing as a result of the adaptation of three, formerly forest-inhabiting species (Lutzomyia intermedia, Lutzomyia whitmani and Lutzomyi migonei) following deforestation. [12] Now, a disease which previously only affected forest workers is becoming more prevalent in urbanisations. Similar increases in risk factors are also driving higher rates of visceral leishmaniasis in north-eastern Brazil. Increased urbanisation, driven by climate change and socio-economic factors, is bringing the adapting sand flies into closer contact with both humans and the domestic canine reservoirs of the disease. [11]

Carrion's disease

Carrion's disease, also known as bartonellosis, is a disease caused by the blood-borne bacteria, Bartonella bacilliformis . The disease is transmitted by the sand fly species Lutzomyia verracarum, as well as lice and fleas, and is found in areas of Peru, Colombia and Ecuador. [13] The lifecycle of the bacteria within Lutzomyia sand flies remain largely unknown, with speculation that the bacteria are spread between sandflies sharing the same breeding grounds and water supplies. The existence of bartonellosis transmission in areas not inhabited by Lutzomyia verracarum suggests that secondary vectors, and potentially other Lutzomyia species, are important in the spread of the disease. [14]

Viruses

A number of arthropod-borne viruses (arboviruses) are transmitted by Lutzomyia sand flies. One such virus of medical importance is the vesicular stomatitis virus (VSV) of the genus Vesiculovirus. Viruses of this genus are typically associated with flu-like symptoms, but have also been known to cause encephalitis. [1] While transmission of the Indiana serotype of VSV by Lutzomyia trapidoi has been demonstrated under laboratory conditions, little is known about the cycles of virus infection between vertebrates and Lutzomyia sand flies. However, they are thought to be supported by both horizontal and transovarial transmission in the sand fly. [15]

Related Research Articles

<i>Leishmania</i> Genus of parasitic flagellate protist

Leishmania is a parasitic protozoan, a single-celled organism of the genus Leishmania that is responsible for the disease leishmaniasis. They are spread by sandflies of the genus Phlebotomus in the Old World, and of the genus Lutzomyia in the New World. At least 93 sandfly species are proven or probable vectors worldwide. Their primary hosts are vertebrates; Leishmania commonly infects hyraxes, canids, rodents, and humans.

<span class="mw-page-title-main">Leishmaniasis</span> Disease caused by parasites of the Leishmania type

Leishmaniasis is a wide array of clinical manifestations caused by parasites of the Trypanosomatida genus Leishmania. It is generally spread through the bite of phlebotomine sandflies, Phlebotomus and Lutzomyia, and occurs most frequently in the tropics and sub-tropics of Africa, Asia, the Americas, and southern Europe. The disease can present in three main ways: cutaneous, mucocutaneous, or visceral. The cutaneous form presents with skin ulcers, while the mucocutaneous form presents with ulcers of the skin, mouth, and nose. The visceral form starts with skin ulcers and later presents with fever, low red blood cell count, and enlarged spleen and liver.

<span class="mw-page-title-main">Sandfly</span> Name of several types of blood-sucking fly

Sandfly or sand fly is a colloquial name for any species or genus of flying, biting, blood-sucking dipteran (fly) encountered in sandy areas. In the United States, sandfly may refer to certain horse flies that are also known as "greenheads", or to members of the family Ceratopogonidae. The bites usually result in a small, intensely itchy bump or welt, the strength of which intensifies over a period of 5-7 days before dissipating. Moderate relief is achieved with varying success through the application of over the counter products such as Benadryl (ingested) or an analgesic cream such as After Bite. Outside the United States, sandfly may refer to members of the subfamily Phlebotominae within the Psychodidae. Biting midges (Ceratopogonidae) are sometimes called sandflies or no-see-ums. New Zealand sandflies are in the genus of sand fly Austrosimulium, a type of black fly.

<i>Phlebotomus</i> Genus of flies

Phlebotomus is a genus of "sand flies" in the Diptera family Psychodidae. In the past, they have sometimes been considered to belong in a separate family, Phlebotomidae, but this alternative classification has not gained wide acceptance.

<span class="mw-page-title-main">Cutaneous leishmaniasis</span> Medical condition

Cutaneous leishmaniasis is the most common form of leishmaniasis affecting humans. It is a skin infection caused by a single-celled parasite that is transmitted by the bite of a phlebotomine sand fly. There are about thirty species of Leishmania that may cause cutaneous leishmaniasis.

<span class="mw-page-title-main">Visceral leishmaniasis</span> Human disease caused by protist parasites

Visceral leishmaniasis (VL), also known as kala-azar or "black fever", is the most severe form of leishmaniasis and, without proper diagnosis and treatment, is associated with high fatality. Leishmaniasis is a disease caused by protozoan parasites of the genus Leishmania.

<span class="mw-page-title-main">Phlebotominae</span> Subfamily of flies

The Phlebotominae are a subfamily of the family Psychodidae. In several countries, their common name is sandfly, but that name is also applied to other flies. The Phlebotominae include many genera of blood-feeding (hematophagous) flies, including the primary vectors of leishmaniasis, bartonellosis and pappataci fever. In the New World, leishmaniasis is spread by sand flies in the genus Lutzomyia, which commonly live in caves, where their main hosts are bats. In the Old World, sand flies in the genus Phlebotomus spread leishmaniasis.

A canine vector-borne disease (CVBD) is one of "a group of globally distributed and rapidly spreading illnesses that are caused by a range of pathogens transmitted by arthropods including ticks, fleas, mosquitoes and phlebotomine sandflies." CVBDs are important in the fields of veterinary medicine, animal welfare, and public health. Some CVBDs are of zoonotic concern.

<i>Leishmania infantum</i> Species of parasitic protist

Leishmania infantum is the causative agent of infantile visceral leishmaniasis in the Mediterranean region and in Latin America, where it has been called Leishmania chagasi. It is also an unusual cause of cutaneous leishmaniasis, which is normally caused by specific lineages. Wild canids and domestic dogs are the natural reservoir of this organism. The sandfly species Lutzomyia longipalpis serves as the primary vector for the transmission of the disease.

<span class="mw-page-title-main">Canine leishmaniasis</span> Disease affecting dogs

Canine leishmaniasis (LEESH-ma-NIGH-ah-sis) is a zoonotic disease caused by Leishmania parasites transmitted by the bite of an infected phlebotomine sandfly. Canine leishmaniasis was first identified in Europe in 1903, and in 1940, 40% of all dogs in Rome were determined to be positive for leishmaniasis. Traditionally thought of as a disease only found near the Mediterranean basin, 2008 research claims new findings are evidence that canine leishmaniasis is currently expanding in continental climate areas of northwestern Italy, far from the recognized disease-endemic areas along the Mediterranean coasts. Cases of leishmaniasis began appearing in North America in 2000, and, as of 2008, Leishmania-positive foxhounds have been reported in 22 U.S. states and two Canadian provinces.

<i>Leishmania donovani</i> Species of intracellular parasite

Leishmania donovani is a species of intracellular parasites belonging to the genus Leishmania, a group of haemoflagellate kinetoplastids that cause the disease leishmaniasis. It is a human blood parasite responsible for visceral leishmaniasis or kala-azar, the most severe form of leishmaniasis. It infects the mononuclear phagocyte system including spleen, liver and bone marrow. Infection is transmitted by species of sandfly belonging to the genus Phlebotomus in Old World and Lutzomyia in New World. The species complex it represents is prevalent throughout tropical and temperate regions including Africa, China, India, Nepal, southern Europe, Russia and South America. The species complex is responsible for thousands of deaths every year and has spread to 88 countries, with 350 million people at constant risk of infection and 0.5 million new cases in a year.

<i>Leishmania tropica</i> Species of protozoan parasite

Leishmania tropica is a flagellate parasite and the cause of anthroponotic cutaneous leishmaniasis in humans. This parasite is restricted to Afro-Eurasia and is a common cause of infection in Afghanistan, Iran, Syria, Yemen, Algeria, Morocco, and northern India.

Leishmania braziliensis is a Leishmania species.

<i>Paleoleishmania</i> Extinct genus of parasitic flagellate protist in the Kinetoplastea class

Paleoleishmania is an extinct genus of kinetoplastids, a monophyletic group of unicellular parasitic flagellate protozoa. At present it is placed in the family Trypanosomatidae. The genus contains two species, Paleoleishmania neotropicum and the type species Paleoleishmania proterus.

Lutzomyia shannoni is a species of fly in the subfamily Phlebotominae, the phlebotomine sand flies. It is native to the Americas from the southeastern United States to northern Argentina. It has a disjunct distribution, and is only found in regions with suitable climates, habitat types, and host animals. It is well known as a vector of the vesicular stomatitis virus, which causes the disease vesicular stomatitis in animals, particularly livestock.

Endotrypanum schaudinni is a parasite found only in Hoffmann's two-toed sloth. This sloth is found throughout South America in forested areas and this parasite is found throughout the population. Hoffman’s two-toed sloth is the only host species so it is of little economic or medical importance in humans. Endotrypanum schaudinni is transferred by an intermediate host of sandflies from the genus Lutzomyia. The highest concentration of flagellates in these insects is found in the pylorus region of this vector which may be important to transmission to the host. E. schaudinni is an intracellular parasite which invades the red blood cells of its definitive host. Little research has been done on the mechanism of action of this or the benefit of residing in the red blood cells but antigenic masking seems a possible reason. E. schaudinni is a hemoflagellate with a size of about 13.5 µm in length and 3.5 µm in diameter. Another point of interest is that E. schaudinni shares many of the same gene sequences as Leishmania, and two unique gene sequences were found in 2009 for diagnostic purposes. This diagnostic technique is an important factor in further studies of this parasite as identifying it but classical biochemical and ultrastructural parameters is unreliable.

<i>Lutzomyia longipalpis</i> Species of fly

Lutzomyia longipalpis is a species complex of sandfly belonging to the family Psychodidae. This species is primarily present in Central and South America, but has also appeared in Mexico. There have been reports of L. longipalpis as far south as Argentina, as they are found in a wide variety of ecological conditions. Both males and females feed on sugars from plants and aphids, but only adult females feed on the blood of other mammals. The species has recently begun appearing in urban areas throughout Brazil, and serves as a key vessel for the propagation of the parasite Leishmania infantum. The presence of these flies appears to be strongly correlated to the presence of domestic chickens in Latin America. The first major urban outbreak of the lethal Visceral leishmanias epidemic was detected in Teresina, Piauí State in the early 1980s following a massive planting of acacias.

<span class="mw-page-title-main">Maria Cristina Ferro</span> Colombian microbiologist

Maria Cristina Ferro de Carrasquilla (1947–2015) was a Colombian microbiologist and Leishmaniasis researcher who worked for more than forty years at the National Health Institute of Colombia. Most of her research was focused on sandflies, which are Leishmaniasis vectors, contributing with the description of three new species: Lutzomyia torvida, Lutzomyia falcata, and Lutzomyia tolimensis. Given her research contributions, a sandfly species was named after her: Lutzomyia ferroae. Ferro also worked with Venezuelan equine encephalitis virus and received the Emeritus Researcher award from the National Health Institute of Colombia in 2007

María Dora Feliciangeli was Professor of Medical Entomology at the University of Carabobo, Venezuela, who worked on the transmission of tropical parasites especially American trypanosomiasis and leishmaniasis. She was Coordinator of the National Reference Center of Sandflies and the Medical Entomology Section.

<i>Phlebotomus papatasi</i>

Phlebotomus papatasi is a species of insects commonly known as sandflies. Due to their ectothermic climate limitations, P. papatasi are confined to regions with temperatures above sixty degrees Fahrenheit for at least three months of the year, spanning over much of the European Mediterranean, North Africa, the Middle East, and Central Asia. Around one thousand sandfly species have been described, belonging to the subfamily Phlebotominae within the family Psychodidae. The Psychodidae family belongs to the order Diptera within the class Insecta of the subphylum Hexapoda of the Arthropod phylum.

References

  1. 1 2 3 4 5 6 7 Young, D. G. & Duncan, M. A. (1994). "Guide to the identification and geographic distribution of Lutzomyia sand flies in Mexico, the West Indies, Central and South America (Diptera: Psychodidae)". Memoirs of the American Entomological Institute (Gainesville) : 1–864.
  2. 1 2 3 Beati, L., Caceres, A. G., Lee, J. A. & Munstermann, L. E. (2004). "Systematic relationships among Lutzomyia sand flies (Diptera : Psychodidae) of Peru and Colombia based on the analysis of 12S and 28S ribosomal DNA sequences". International Journal for Parasitology . 34 (2): 225–234. doi:10.1016/j.ijpara.2003.10.012. PMID   15037108.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  3. ↑ Poinar, G. (2008). "Lutzomyia adiketis sp. n. (Diptera: Phlebotomidae), a vector of Paleoleishmania neotropicum sp. n. (Kinetoplastida: Trypanosomatidae) in Dominican amber". Parasites & Vectors. 1 (1): 22. doi: 10.1186/1756-3305-1-22 . PMC   2491605 . PMID   18627624.
  4. ↑ Andrade, J. D. & Brazil, R. P. (2003). "Relationships of New World phlebotomine sand flies (Diptera : Psychodidae) based on fossil evidence". Memórias do Instituto Oswaldo Cruz . 98: 145–149. doi: 10.1590/s0074-02762003000900022 . PMID   12687775.
  5. 1 2 Killick-Kendrick, R. (1999). "The biology and control of phlebotomine sand flies". Clinics in Dermatology . 17 (3): 279–289. doi:10.1016/S0738-081X(99)00046-2. PMID   10384867.
  6. ↑ Grimaldi, G. & Tesh, R. B. (1993). "Leishmaniases of the New-World - Current Concepts and Implications for Future Research". Clinical Microbiology Reviews . 6 (3): 230–250. doi:10.1128/CMR.6.3.230. PMC   358284 . PMID   8358705.
  7. 1 2 3 Gramiccia, M. & Gradoni, L. (2005). "The current status of zoonotic leishmaniases and approaches to disease control". International Journal for Parasitology . 35 (11–12): 1169–1180. doi:10.1016/j.ijpara.2005.07.001. PMID   16162348.
  8. ↑ Alexander, Bruce; Lopes de Carvalho, Renata; McCallum, Hamish; Pereira, Marcos Horácio (December 2002). "Role of the Domestic Chicken (Gallus gallus)in the Epidemiology of Urban Visceral Leishmaniasis in Brazil". Emerging Infectious Diseases. 8 (12): 1480–1485. doi:10.3201/eid0812.010485. PMC 2738513. PMID 12498667.
  9. ↑ Croan, D. G., Morrison, D. A. & Ellis, J. T. (1997). "Evolution of the genus Leishmania revealed by comparison of DNA and RNA polymerase gene sequences". Molecular and Biochemical Parasitology . 89 (2): 149–159. doi:10.1016/S0166-6851(97)00111-4. PMID   9364962.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  10. ↑ Bates, P. A. (2007). "Transmission of Leishmania metacyclic promastigotes by phlebotomine sand flies". International Journal for Parasitology . 37 (10): 1097–1106. doi:10.1016/j.ijpara.2007.04.003. PMC   2675784 . PMID   17517415.
  11. 1 2 Desjeux, P. (2001). "The increase in risk factors for leishmaniasis worldwide". Transactions of the Royal Society of Tropical Medicine and Hygiene . 95 (3): 239–243. doi: 10.1016/S0035-9203(01)90223-8 . PMID   11490989.
  12. ↑ Peterson, A. T. & Shaw, J. (2003). "Lutzomyia vectors for cutaneous leishmaniasis in Southern Brazil: ecological niche models, predicted geographic distributions, and climate change effects". International Journal for Parasitology . 33 (9): 919–931. doi:10.1016/S0020-7519(03)00094-8. PMID   12906876.
  13. ↑ Maguina, C., Garcia, P. J., Gotuzzo, E., Cordero, L. & Spach, D. H. (2001). "Bartonellosis (Carrion's disease) in the modern era". Clinical Infectious Diseases . 33 (6): 772–779. doi:10.1086/322614. PMID   11512081. S2CID   16680459.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  14. ↑ Billeter, S. A., Levy, M. G., Chomel, B. B. & Breitschwerdt, E. B. (2008). "Vector transmission of Bartonella species with emphasis on the potential for tick transmission". Medical and Veterinary Entomology . 22 (1): 1–15. doi: 10.1111/j.1365-2915.2008.00713.x . PMID   18380649.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  15. ↑ Tesh, R. B., Chanioti, B. N. & Johnson, K. M. (1971). "Vesicular Stomatitis Virus, Indiana Serotype - Multiplication in and Transmission by Experimentally Infected Phlebotomine Sandflies (Lutzomyia-Trapidoi)". American Journal of Epidemiology . 93 (6): 491–&. doi:10.1093/oxfordjournals.aje.a121284. PMID   4327378.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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