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The members of the Triatominae , a subfamily of the
Reduviidae The Reduviidae are a large cosmopolitan family of the order Hemiptera (true bugs). Among the Hemiptera and together with the Nabidae almost all species are terrestrial ambush predators: most other predatory Hemiptera are aquatic. The main exampl ...
, are also known as conenose bugs, kissing bugs (so-called from their habit of feeding from around the mouths of people), or vampire bugs. Other local names for them used in The Americas include ''barbeiros'', ''vinchucas'', ''pitos'', ''chipos'' and ''chinches''. Most of the 130 or more species of this subfamily feed on vertebrate blood; a very few species feed on invertebrates. They are mainly found and widespread in the Americas, with a few species present in Asia and Africa. These bugs usually share shelter with nesting vertebrates, from which they suck blood. In areas where
Chagas disease Chagas disease, also known as American trypanosomiasis, is a tropical parasitic disease caused by ''Trypanosoma cruzi''. It is spread mostly by insects in the subfamily '' Triatominae'', known as "kissing bugs". The symptoms change over the co ...
occurs (from the southern
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 Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
to northern
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 List of South American countries by area, second-largest ...
), all triatomine species are potential
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 ...
s of the Chagas disease parasite ''
Trypanosoma cruzi ''Trypanosoma cruzi'' is a species of parasitic euglenoids. Among the protozoa, the trypanosomes characteristically bore tissue in another organism and feed on blood (primarily) and also lymph. This behaviour causes disease or the likelihood o ...
'', but only those species that are well adapted to living with humans (such as ''
Triatoma infestans ''Triatoma infestans'', commonly called winchuka or vinchuca in Argentina, Bolivia, Uruguay and Chile, barbeiro in Brazil, chipo in Venezuela and also known as "kissing bug" or "barber bug" in English, is a blood-sucking bug (like virtually a ...
'' and ''
Rhodnius prolixus ''Rhodnius prolixus'' is the principal triatomine vector of the Chagas parasite due to both its sylvatic and domestic populations in northern South America as well as to its exclusively domestic populations in Central America. It has a wide ra ...
'') are considered important vectors. Also, proteins released from their bites have been known to induce
anaphylaxis Anaphylaxis is a serious, potentially fatal allergic reaction and medical emergency that is rapid in onset and requires immediate medical attention regardless of use of emergency medication on site. It typically causes more than one of the follo ...
in sensitive and sensitized individuals.


History

At the beginning of the 19th century,
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
made one of the first reports of the existence of triatomines in America in his ''Journal and Remarks'', published in 1839 and commonly known as '' The Voyage of the Beagle''. The following is an extract which he based on his journal entry dated 26 March 1835: Considerable medical speculation has occurred as to whether or not Darwin's contact with triatomines in Argentina was related to his later bouts of long-term illness, though it is unlikely to have been caused on this specific occasion, as he made no mention of the fever that usually follows the first infection.


Discovery of triatominae's relation with Chagas disease

In 1909, Brazilian doctor
Carlos Chagas Carlos Justiniano Ribeiro Chagas, or Carlos Chagas (; July 9, 1879 – November 8, 1934), was a Brazilian sanitary physician, scientist, and bacteriologist who worked as a clinician and researcher. He discovered Chagas disease, also called ''A ...
discovered that these insects were responsible for the transmission of ''T. cruzi'' to many of his patients in Lassance, a village located on the banks of the
São Francisco River The São Francisco River (, ) is a large river in Brazil. With a length of , it is the longest river that runs entirely in Brazilian territory, and the fourth longest in South America and overall in Brazil (after the Amazon, the Paraná and t ...
in
Minas Gerais Minas Gerais () is a state in Southeastern Brazil. It ranks as the second most populous, the third by gross domestic product (GDP), and the fourth largest by area in the country. The state's capital and largest city, Belo Horizonte (literall ...
(
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
). Poor people living there complained of some insects they called ''barbeiros'' that bite during the night. Carlos Chagas put his first observations in words: Another Brazilian, Herman Lent, former student of Carlos Chagas, became devoted to the research of the triatomines and together with Peter Wygodzinsky made a revision of the Triatominae, a summary of 40 years of studies on the triatomines up to 1989.


Biological aspects


Lifecycle

Triatomines undergo
incomplete metamorphosis Hemimetabolism or hemimetaboly, also called incomplete metamorphosis and paurometabolism,McGavin, George C. ''Essential Entomology: An Order-by-Order Introduction''. Oxford: Oxford University Press, 2001. pp. 20. is the mode of development of certa ...
. A wingless first-
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 ...
nymph hatches from an egg, and may be small as 2 mm. It passes successively through second, third, fourth, and fifth instars. Finally, the fifth instar turns into an adult, acquiring two pairs of wings.


Ecology

All triatomine
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...
instars and adults are
haematophagous 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 ...
and require the stability of a sheltered environment, where they aggregate. Most species are associated with wild, nesting
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 ...
s and are named " sylvatic" triatomines. These live in ground burrows with
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s or
armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, alo ...
s, or in tree dwellings with bats, birds,
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their l ...
s, or
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered No ...
s. Few species (5%) live in human dwellings or in the surroundings of human houses (peridomicile) in the shelters of domestic animals, these are named " domestic" species. Many sylvatic species are in process of domiciliation (i.e. "semidomestic").


Behavior

Most triatomines aggregate in refuges during day and search for blood during night, when the host is asleep and the air is cooler. Odors and heat guide these insects to their hosts. Carbon dioxide emanating from breath, as well as ammonia, short-chain amines, and carboxylic acids from skin, hair, and exocrine glands from vertebrate animals, are among the volatiles that attract triatomines. Vision also serves triatomines for orientation. At night, adults of diverse species fly to houses attracted by light. Adults produce a pungent odor (
isobutyric acid Isobutyric acid, also known as 2-methylpropanoic acid or isobutanoic acid, is a carboxylic acid with structural formula ( CH3)2CH COOH. It is an isomer of ''n''-butyric acid. It is classified as a short-chain fatty acid. Deprotonation or esterif ...
) when disturbed, and are also capable of producing a particular sound by rubbing the rostrum over a stridulatory sulcus under its head (
stridulation Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
).


Epidemiology

Domestic and sylvatic species can carry the Chagas parasite to humans and wild mammals; birds are immune to the parasite. ''T. cruzi'' transmission is carried mainly from human to human by domestic kissing bugs; from the vertebrate to the bug by blood, and from the bug to the vertebrate by the insect's feces, and not by its saliva as occurs in most bloodsucking arthropod vectors such as malaria mosquitoes. Triatomine infestation especially affects older dwellings. One can recognize the presence of triatomines in a house by its feces, exuviae, eggs, and adults. Triatomines characteristically leave two kinds of
feces Feces ( or faeces), known colloquially and in slang as poo and poop, 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 large intestine. Feces contain a rela ...
like strikes on walls of infected houses; one is white with
uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the metabolic breakdown ...
, and the other is dark (black) containing
heme Heme, or haem (pronounced / hi:m/ ), is a precursor to hemoglobin, which is necessary to bind oxygen in the bloodstream. Heme is biosynthesized in both the bone marrow and the liver. In biochemical terms, heme is a coordination complex "consis ...
. Whitish or pinkish eggs can be seen in wall crevices. After having had a blood meal, the insects sometimes show a limited mobility and can be identified easily.


Controlling triatomine infestations


Insecticide treatment

Synthetic
pyrethroids A pyrethroid is an organic compound similar to the natural pyrethrins, which are produced by the flowers of pyrethrums (''Chrysanthemum cinerariaefolium'' and '' C. coccineum''). Pyrethroids are used as commercial and household insecticides. ...
are the main class of
insecticide Insecticides are substances used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Insecticides are used in agriculture, medicine, industry and by consumers. Insecticides are claimed t ...
s used to control triatominae infestations. Insecticide treatment is more effective on nonporous surfaces, such as hardwood timber, fired bricks, and plastered walls, than on porous surfaces such as mud. A single treatment with insecticide typically protects against triatomine infestation for a year or more on timber walls vs. 2–3 months on adobe walls. Wettable powders, suspension concentrates, and insecticide paints can improve treatment effectiveness on porous surfaces. Rates of
insecticide resistance Pesticide resistance describes the decreased susceptibility of a pest population to a pesticide that was previously effective at controlling the pest. Pest species evolve pesticide resistance via natural selection: the most resistant specimens su ...
among triatomines are fairly low due to their long lifecycle and low genetic variability, but some instances of resistance have been reported, particularly among''
Triatoma infestans ''Triatoma infestans'', commonly called winchuka or vinchuca in Argentina, Bolivia, Uruguay and Chile, barbeiro in Brazil, chipo in Venezuela and also known as "kissing bug" or "barber bug" in English, is a blood-sucking bug (like virtually a ...
'' populations in Bolivia and Argentina.


Tribes, genera, and numbers of described species

The monophyletic nature of the Triatominae subfamily is strongly supported by molecular data, indicating that their blood-sucking character has occurred only once within the
Reduviidae The Reduviidae are a large cosmopolitan family of the order Hemiptera (true bugs). Among the Hemiptera and together with the Nabidae almost all species are terrestrial ambush predators: most other predatory Hemiptera are aquatic. The main exampl ...
. The classification within the subfamily is not stable, with different proposed relationships among the tribes and genera. The classification below largely follows Galvão et al. 2003, but in 2009 this same author eliminated the tribe Linshcosteini and also eliminated the genera '' Meccus'', ''
Mepraia ''Mepraia'' is a genus in the subfamily Triatominae, endemic in Chile, and vectors of Chagas disease Chagas disease, also known as American trypanosomiasis, is a tropical parasitic disease caused by ''Trypanosoma cruzi''. It is spread mostl ...
'', and '' Nesotriatoma''Schofield CJ, Galvão C. (2009) Classification, evolution, and species groups within the Triatominae. Acta Trop. 110: 88-100. * †'' Paleotriatoma''
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar,
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
(
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...
) ;Alberproseniini (monotypic) * '' Alberprosenia'' 2


Bolboderini

# '' Belminus'' 9 # '' Bolbodera'' 1 # '' Microtriatoma'' 2 # '' Parabelminus'' 2 ;Cavernicolini (monotypic) * '' Cavernicola'' 2 ; Linshcosteini # ''
Linshcosteus ''Linshcosteus'' is a genus of assassin bugs in the subfamily Triatominae (the kissing bugs). It is the only genus of Triatomines restricted to the Old World within the mostly Neotropical subfamily Triatominae (a few '' Triatoma'' species are kno ...
'' Distant, 1904 (6)


Rhodniini

# '' Psammolestes'' 3 # ''
Rhodnius ''Rhodnius'' is a genus of assassin bugs in the subfamily Triatominae (the kissing bugs), and is an important vector in the spread of Chagas disease. The ''Rhodnius'' species were important models for Sir Vincent Wigglesworth's studies of insect ...
'' 16


Triatomini

# '' Dipetalogaster'' 1 # '' Eratyrus'' 2 # '' Hermanlentia'' 1 # '' Meccus'' 6 # ''
Mepraia ''Mepraia'' is a genus in the subfamily Triatominae, endemic in Chile, and vectors of Chagas disease Chagas disease, also known as American trypanosomiasis, is a tropical parasitic disease caused by ''Trypanosoma cruzi''. It is spread mostl ...
'' 2 # '' Nesotriatoma'' 3 # '' Panstrongylus'' 13 # '' Paratriatoma'' 1 # ''
Triatoma ''Triatoma'' is a genus of assassin bug in the subfamily Triatominae (kissing bugs). The members of ''Triatoma'' (like all members of Triatominae) are blood-sucking insects that can transmit serious diseases, such as Chagas disease. Their saliva ...
'' 67


Most important vectors

All 138 Triatominae species are potentially able to transmit ''T. cruzi'' to humans, but these five species are the most epidemiologically important vectors of Chagas disease. * ''
Triatoma infestans ''Triatoma infestans'', commonly called winchuka or vinchuca in Argentina, Bolivia, Uruguay and Chile, barbeiro in Brazil, chipo in Venezuela and also known as "kissing bug" or "barber bug" in English, is a blood-sucking bug (like virtually a ...
'' * ''
Rhodnius prolixus ''Rhodnius prolixus'' is the principal triatomine vector of the Chagas parasite due to both its sylvatic and domestic populations in northern South America as well as to its exclusively domestic populations in Central America. It has a wide ra ...
'' * '' Triatoma dimidiata'' * ''
Triatoma brasiliensis ''Triatoma brasiliensis'' is now considered the most important Chagas disease vector in the semiarid areas of northeastern Brazil. ''T. brasiliensis'' occurs in 12 Brazilian states, including Maranhão, Piauí, Ceará, Rio Grande do Norte, and P ...
'' * ''
Panstrongylus megistus ''Panstrongylus megistus'' is a blood-drinking insect in the subfamily Triatominae. It is found in the Guianas, Brazil, Bolivia, Paraguay, Uruguay and Argentina, and is an important vector of ''Trypanosoma cruzi'' (the causative agent of Chagas ...
''


See also

* Hematophagy


References


Further reading

* Brenner RR, Stoka AM (1987) ''Chagas’ disease vectors''. I, II and III. CRC Press. Boca Ratón * Dujardin JP, Schofield CJ, Panzera F (2000) "Les vecteurs de la maladie de Chagas: recherches taxonomiques, biologiques et génétiques". ''Academie Royale des Sciences d'Ultre-Mer''. Belgium.


External links


The Kiss of Death: Chagas' Disease in the Americas

Taxonomy and general information on Triatominae bugs from metapathogen.com

World Health Organisation guide to Triatominae
* {{Taxonbar, from=Q13381446 Insect vectors of human pathogens Reduviidae Hemiptera subfamilies Hematophages