Biology, ecology, and systematics of Triatominae ... · named the species Trypanosoma cruzi. He...

Historical background

The first triatomine bug species was de-scribed scientifically by Carl DE GEER

(1773), (Fig. 1), but according to LENT &WYGODZINSKY (1979), the first report on as-pects and habits dated back to 1590, byReginaldo de Lizárraga. While travelling toinspect convents in Peru and Chile, thispriest noticed the presence of largehematophagous insects that attacked atnight. In subsequent reports, various travel-ers and naturalists also mentioned the pres-ence of these insects in South America. Oneof the most celebrated reports was byCharles Darwin, during his trip to SouthAmerica on the H.M.S. Beagle in 1835,when he wrote: “One which I caught atIquique, (for they are found in Chile andPeru) was very empty. When placed on atable, and though surrounded by people, if afinger was presented, the bold insect wouldimmediately protrude its sucker, make acharge, and if allowed, draw blood. No painwas caused by the wound. It was curious towatch its body during the act of sucking, asin less than ten minutes it changed from be-ing as flat as a wafer to a globular form...“

(DARWIN 1871; LENT & WYGODZINSKY

1979).

American trypanosomiasis or Chagasdisease was discovered in 1909 under curi-ous circumstances. In 1907, the Brazilianphysician Carlos Ribeiro Justiniano dasChagas (1879-1934) was sent by OswaldoCruz to Lassance, a small village in the stateof Minas Gerais, Brazil, to conduct an anti-malaria campaign in the region where a rail-way was being built. The young doctor (28years old) stayed in the area for about oneyear, during which time a railroad engineer,Cantarino Mota, alerted him to the pres-ence of hematophagous insects referred tolocally as “barbeiros“ (the local term for tri-atomines) (Fig. 2). Alerted to the presenceof these insects inside the human dwellings,the doctor decided to investigate the possi-bility that they might transmit some parasiteto humans, since besides malaria, he had de-tected clinical signs that were difficult to in-terpret. The local population complainedabout an uncomfortable feeling referred toas “baticum“ (a folk term for palpitation, lit-erally meaning “drumming“ or “pounding“),and Chagas observed arrhythmia and signs

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Biology, ecology, and systematics ofTriatominae (Heteroptera, Reduviidae),

vectors of Chagas disease, and implications for human health1

J . J U R B E R G & C . G A L V Ã O

Abstract: The members of the subfamily Triatominae (Heteroptera, Reduviidae) are vectors of Try-panosoma cruzi (CHAGAS 1909), the causative agent of Chagas disease or American trypanosomiasis. Asimportant vectors, triatomine bugs have attracted ongoing attention, and, thus, various aspects of theirsystematics, biology, ecology, biogeography, and evolution have been studied for decades. In the presentpaper the authors summarize the current knowledge on the biology, ecology, and systematics of thesevectors and discuss the implications for human health.

Key words: Chagas disease, Hemiptera, Triatominae, Trypanosoma cruzi, vectors.

Denisia 19, zugleich Kataloge der OÖ. Landesmuseen

Neue Serie 50 (2006), 1096–11161 This paper is dedicated to Ernst Heiss on the occasion of his 70th birthday.

© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

of heart failure among local residents, alongwith reports of unexplained sudden death.Upon dissecting the triatomines he foundflagellates in their intestinal tract. Believingthat they belonged to Trypanosoma mi-nasense, which infected black tufted-earmarmosets (Callithrix penicillata) in the sameregion, he sent several triatomine specimensto Oswaldo Cruz in Rio de Janeiro to feedon uninfected marmosets. Several weeks lat-er he returned to Rio de Janeiro to discovera new trypanosome in the blood of one ofthe animals. As a tribute to his mentor henamed the species Trypanosoma cruzi. Hethen returned to Minas Gerais to attempt toidentify the parasite’s vertebrate host. Afternumerous negative blood samples, he foundan infected cat. Some 30 days later he re-turned to the house where he had discov-ered the infected animal and found a littlegirl named Berenice, just two years old, whowas febrile and presented circulating formsof T. cruzi in her bloodstream.

From 1909 to 1912, Chagas described anew disease, its causative agent, naturalreservoirs, and the vector, a fact unparal-leled in international medicine to this dayand a milestone in the history of medicine,since the discovery was made in the inverseorder as compared to the usual (where thediscovery of a disease generally precedesthat of its causative agent). As a result of hiswork, in 1912 Chagas received theSchaudinn Award from the Institute ofTropical Diseases in Hamburg, Germany(CHAGAS FILHO 1968). Thus, one and thesame researcher, in inverse order and in ashort space of time, discovered a new diseasethat would later bear his last name, first rec-ognizing the vector, next the parasite andreservoirs, and finally the clinical disease inhumans (CHAGAS 1909). Chagas’ discoverywas overlooked by the Brazilian scientificcommunity, as represented by the NationalAcademy of Medicine, and was treated withdisbelief for more than 20 years, becausesome scientists questioned the very exist-ence of the disease. It was in Argentina in1935 that Salvador Mazza submitted studieson the disease to the Annual Meeting of theArgentine Society of Tropical Medicineand, together with Cecílio Romaña, gave anew dimension and credibility to the prob-lem (CHAGAS FILHO 1968).

For more than one century, since thefirst description by De Geer, triatomineswere studied merely from a descriptive pointof view. However, beginning in 1909, whenChagas discovered the disease, and due itsnewly acquired relevance to human health,studies began on the clinical form of the dis-ease, the protozoan, and the vertebratehosts, as well as the vector biology andtransmission mechanisms. Advances in vec-tor taxonomy began with Arthur Neiva, oneof the most important scientists in thisphase, who, in 1911, began describing vari-ous species, culminating with the publica-tion of his dissertation “Revisão do gêneroTriatoma Lap.“ in 1914. Important mono-graphs were published subsequently by PIN-TO (1925) and DEL PONTE (1930), in addi-tion to other extensive studies published byNEIVA & LENT (1936, 1941), USINGER

(1944), ABALOS & WYGODZINSKY (1951),and RYCKMAN (1962), culminating in thegrand work by LENT & WYGODZISNKY

(1979).

Trypanosoma cruzi and Chagas disease

Chagas disease or American trypanoso-miasis was primarily a zoonosis, a parasiticdisease of wild animals transmitted by syl-vatic species of triatomine bugs. The adap-tation of some triatomine species to humandwellings was secondary, as was the para-site’s domiciliary cycle. American try-panosomiasis is now an endemic disease af-fecting mostly Latin America, primarily inrural populations of Central and SouthAmerica, where it is an important publichealth problem. According to World HealthOrganization, an estimated 100 million peo-ple are at risk of the disease, with 18 millioncurrently infected (DIAS & COURA 1997).Currently there is no vaccine or effectivecure for chronic Chagas disease.

Trypanosoma cruzi, the etiological agentof the disease, is a flagellate protozoan of theorder Kinetoplastida, family Trypanosomati-dae. It is some 20-thousandths of a millime-ter in length and has an elongated bodywith an undulating membrane allowing itsmovement in the bloodstream.

Unlike other diseases transmitted byhematophagous insects, in this case infec-tion does not occur through the insect’s sali-

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Fig. 1: Male of Triatoma rubrofasciata (DE

GEER 1773), the first species described.


va. When the triatomine bug bites, it defe-cates during or right after bloodsucking,eliminating infective forms of the parasite inits feces. These forms can penetrate activelyeither through the bite’s orifice, the mucosa,or small wounds and scratches on the skin.Vector-borne transmission is the most im-portant form of transmission, but it can alsotake place through transfusion, transplacen-tally, through organ transplants, accidentalingestion of triatomines, laboratory acci-dents, or even inadequate handling of in-fected animal carcasses.

Animals infected with T. cruzi are al-ways mammals, as the parasite cannot de-velop in the blood of birds, reptiles, or am-phibians. A triatomine bug that has suckedthe blood of a mammal (including a human)infected with T. cruzi acquires the infectionand the protozoan then reproduces, multi-plying in the insect’s digestive tract and pro-ducing the infective forms which are ex-pelled in the feces. Infection remains in theinsect throughout its lifespan and can occurboth in the nymphs and adults. If the insectsucks infected blood early in life (1st instarnymph), it acquires the infection and willremain infected throughout its life. Onlythe eggs are not affected, so the second gen-eration remains uninfected until its first in-gestion of infected blood (i.e. there is notransovarian transmission). Therefore tri-atomine bugs reared in the laboratory withblood from uninfected animals can be usedsafely in experiments. After biting and suck-ing the blood of humans or animals, the sa-tiated triatomine bug defecates close to thesite of the bite, and these feces can containthe infective form of T. cruzi. The parasitesquickly penetrate into the bloodstreamthrough the triatomine’s bite, some minorwound, or through the mucosa (of the eyes,nose, or mouth). Upon entering the blood-stream the parasites are transported to themuscles or organs (mainly the heart and di-gestive tract), where they multiply andcause lesions. Penetration of Trypanosomacruzi through the skin can cause a local re-action known as a chagoma, and anotherimportant sign is unilateral bipalpebralswelling, leaving the patient’s eye practical-ly closed. This is the so-called Romaña’ssign, named after its discoverer, an Argen-tine physician.

There are three distinct phases in thedisease: acute, indeterminate, and chronic.In the acute phase (3-4 weeks), the infec-tion varies from an asymptomatic to a severeand fatal form, the latter mainly in childrenor debilitated individuals, characterized byhigh fever, while other symptoms like diar-rhea and vomiting can appear when the di-gestive tract is affected. The indeterminatephase is characterized by low parasitemiawithout clinical signs, which can persist orevolve into a chronic disease. The chronicform normally appears 10 to 15 years afterthe acute phase, and Chagasic cardiopathyis the most common manifestation, the di-gestive form producing visceromegalies, es-pecially megaesophagus and megacolon.

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Fig. 2: Panstrongylus megistus (BURMEISTER

1835), the triatomine species in whichChagas found Trypanosoma cruzi.


Chagas disease vectors

Members of the Triatominae arehemimetabolous insects whose developmentfrom egg to adult includes five nymphalstages (Fig. 3). They are obligatoryhematophagous insects in all phases of theirdevelopment. The amount of blood ingestedvaries according to the species as well as inrelation to the stage, and generally the 4th

and 5th instar nymphs are the ones that feedthe most. In general most species are noc-turnal, and during the day they remain intheir resting places, although they maysometimes go out to suck blood during theday under adverse conditions. In coloniesreared in the laboratory, the bugs seek foodsources in broad daylight.

When they finish a meal their bodychanges appearance, since the volume ofblood ingested is so great that the abdomendilates considerably, giving them a globoseappearance by stretching the intersegmentaland connexival membranes. Some speciesmay defecate while feeding, while othersdefecate soon afterwards or even abandonthe food source and defecate far from thesucking site. This fact characterizes the var-ious species as either good or bad transmit-ters of the disease, as the infective forms ofT. cruzi are expelled in the feces of the in-fected bugs.

These insects generally suck the bloodof their victims at night, while they areasleep. The bites generally occur on uncov-ered parts of the body. That is why the pop-ular name for the triatomine bug in Brazil is“barbeiro“ (barbeiro being “barber“ in Por-tuguese). The sleeping persons are unawareof the bugs, because the bites are generallypainless due to the anesthetic and anti-clot-ting action of the saliva. Cases of hypersen-sitivity can occur, but are rare.

Triatomines are vector insects and para-sites with slow development, whose life-cy-cle can range from 3-4 months for Rhodniusprolixus STÅL up to two years for Panstrongy-lus megistus (BURMEISTER). They live in pro-tected environments, the resting places oftheir hosts, which can be either sylvatic ordomestic animals or humans. Of the knownspecies, about a dozen are epidemiologicallyimportant, since they have become domicil-iated due to manmade disequilibria in na-ture (LENT & WYGODZINSKY 1979).

Triatominae external morphology.Most triatomine species can be identified onthe basis of their external morphology. Thegeneral appearance of triatomines is similarto that of other reduviids, and the generalmorphology is shown in Fig. 4. Detailedschemata of morphological traits were pub-lished by LENT & WYGODZINSKY (1979) andseveral scanning electron microscopy illus-trations were provided by CARCAVALLO etal. (1998/99). Adults differ from nymphs bythe presence of ocelli, and well-developedexternal genitalia and wings (with the ex-ception of two species of Mepraia, which dis-play wing polymorphism – see JURBERG et al.2002). Females have a appointed or trun-

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Fig. 3: Eggs, nymphs and female ofPanstrongylus megistus (BURMEISTER 1835).


cate abdominal apex, while in males theapex is rounded. Females are larger thanmales, and the largest triatomine is Dipetalo-gaster maxima (UHLER), which can reach 44mm in length. The smallest species is Alber-prosenia goyovargasi MARTÍNEZ & CARCAV-ALLO, 5 mm long. The length of mostspecies usually varies from 20 to 28 mm. Thecolor pattern (Figs 5-10) varies, with anoverall black or piceous color and spottedpatterns of yellow, brown, orange, or red(JURBERG et al. 2004). The morphology oftriatomine eggs and nymphs (Fig. 11) hasbeen the target of several studies, and a sum-mary of these works was provided byGALVÃO et al. (2005).

Systematics and current classification

Current classification of this subfamilyis based mainly on the revision by LENT &WYGODZINSKY (1979), the most importantsystematic work on this subfamily. Sincethat revision there has been considerablework, including the descriptions of severalnew taxa, and therefore in the present paperthe authors provide a summary of the recentclassification updated from GALVÃO (2003),GALVÃO et al. (2003) and FORERO et al.(2004) (see Table 1). Although generallyrecognized as a monophyletic subfamily ofthe Reduviidae adapted to a blood-feedingstrategy (USINGER 1944; LENT & WYGO-DZINSKY 1979; CLAYTON 1990; SCHUH &SLATER 1995), the subfamily Triatominaestill has several serious phylogenetic ques-tions in regard to its origin and evolution(SCHAEFER 1998, 2003, 2005). SCHOFIELD

(1988) proposed a concept of polyphyleticorigin of the triatomines, and according tohis view the Asiatic fauna consists of at leasttwo independent lineages derived from dif-ferent reduviid stocks. The first Asiatic lin-eage consists of several species of Triatoma,which had evolved from what was originallythe New World species Triatoma rubrofascia-ta (DE GEER) after its introduction into theOld World. The other lineage is representedby the genus Linshcosteus, a supposedly au-tochthonous Asiatic lineage of blood-feed-ing reduviids. This view was supported byGORLA et al. (1997) through morphometricanalysis, where all species of Linshcosteuswere shown to be unrelated to the species ofTriatoma recorded from the Old World. On

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Fig. 4: Triatominae general externalmorphology.

Fig. 5: Color pattern of triatomine bugs: Rhodnius stali LENT, JURBERG & GALVÃO 1993.


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Fig. 6: Color pattern of triatomine bugs: Triatoma rubrovaria (BLANCHARD

1843).

Fig. 7: Color pattern of triatomine bugs: Triatoma sherlocki PAPA, JURBERG,CARCAVALLO, CERQUEIRA & BARATA 2002.

Fig. 8: Color pattern of triatomine bugs: Triatomadimidiata (LATREILLE 1811).

Fig. 9: Color pattern of triatomine bugs: Triatomainfestans (KLUG 1834).


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the other hand, molecular cladistic analysisof 57 species (HYPSA et al. 2002) showed thegenus Linshcosteus and Triatoma rubrofasciataas sister groups. The Linshcosteus-T. rubro-fasciata clade nests firmly within Triatomini,supporting a monophyletic origin of Tri-atominae. According to SCHAEFER (2005)the most important question about phyloge-netic relationships of the Triatominae canbe divided in two parts: Is Triatominae actu-ally a truly independent subfamily at all?And, if Triatominae is a separate subfamily,to which other reduviid subfamilies is Tri-atominae closely related? Attempting tosolve this puzzle, several scientists have usedmolecular data to infer phylogenetic rela-tionships (GARCIA & POWELL 1998;STOTHARD et al. 1998; LYMAN et al. 1999;GARCIA 1999; MARCILLA et al. 2002; SAINZ

et al. 2004). Unfortunately, most of thesepapers are based on limited taxa samplingand are hence unable to solve the questionsof the (entire) Triatomine subfamily phy-logeny. A recently published paper (PAULA

et al. 2005) analyzes mitochondrial rDNAsequences of several triatomine genera andsome representatives of other reduviid sub-families. This analysis suggests that the Tri-atominae itself and Rhodnius and Triatomagenera are polyphyletic. Although this pa-per is an important contribution, it must besupported by additional analyses includingmorphological characters. In conclusion, allthese studies indicate that we still have in-sufficient knowledge to answer these ques-tions and that much additional research isneeded.

Extensive dichotomous keys for theidentification of Triatominae species wereprovided by LENT & WYGODZINSKY (1979),LENT et al. (1994) and CARCAVALLO et al.(1998/99).

Geographical distribution

The vast majority of Triatominae speciesare found only in the New World; a fewspecies are found in East Asia and on thecoast of Australia. In the Neotropical andNearctic regions the species are found be-tween 42°N and 46°S. One species, Tri-atoma rubrofasciata is tropicopolitan, and anadditional seven species of Triatoma occur insouthern and southeastern Asia and north-

Subfamily Tribes Genera Number of valid names

Triatominae Alberproseniini Alberprosenia 2Bolboderini Belminus 6

Bolbodera 1Microtriatoma 2Parabelminus 2

Cavernicolini Cavernicola 2Linshcosteini Linshcosteus 6Rhodniini Psammolestes 3

Rhodnius 16Triatomini Dipetalogaster 1

Eratyrus 2Hermanlentia 1Meccus 6Mepraia 2Nesotriatoma 3Panstrongylus 13Paratriatoma 1Triatoma 67

Total 18 136

Tab. 1: Current systematic classification of the Triatominae subfamily(updated from GALVÃO 2003; GALVÃO et al. 2003; FORERO et al. 2004).

Fig. 10: Color pattern oftriatomine bugs: Triatomanitida USINGER 1939.


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ern Australia. Linshcosteus (Fig. 12) is theonly genus restricted to the Old World,specifically, to the Indian subcontinent(GALVÃO et al. 2003). CARCAVALLO et al.(1999) provided several maps showing thegeographical distribution and altitudinal/latitudinal dispersion of all American Tri-atominae species.

Habitats

Knowledge of triatomine species biologyin their natural habitats is scarce, and forseveral species the natural ecotopes havenot been described. Triatominae species arefound in almost any habitat offering a de-gree of permanence, climatic shelter, andaccess to a blood source. Most species toler-ate a range of air humidity between 30-80%, and temperatures of 24-28 °C are satis-factory to them. Their development is usu-ally slow at temperatures below 16 °C,whereas temperatures above 40 °C arelethal. Several species have specific habi-tats. For example, Triatoma infestans is foundalmost exclusively in human dwellings,Cavernicola lenti, a sylvatic species, is closelyrelated to tree hollows inhabited by bats,and species from of the genus Psammolestesoccur only in the nests of certain birds. Mosttriatomine species are sylvatic, living in birdnests, animal dens, under tree bark or in treehollows, bromeliads, palm trees, and otherecotopes, feeding on various animals.GAUNT & MILES (2000) summarize thehabitats of the triatomine genera Rhodniusand Triatoma, showing that most species inthe former genus live in or associated withpalm trees, while most species in the lattergenus live in or associated with rocky/terres-trial habitats. However, during their evolu-tionary process some species acquired thecapacity to colonize manmade structuresnear human dwellings, like chicken coops,pigsties and such species are thus referred toas peridomiciliated. Others can colonize theinterior of human dwellings and are thuscalled domiciliated. The latter are more im-portant, since they account for transmissionof the disease to humans (CARCAVALLO etal. 1998/99). Many rural inhabitants live indwellings with straw roofs and mud walls,and these provide various forms of shelterfor the triatomines (Fig. 13).

Species and author CountriesAlberprosenia goyovargasi MARTÍNEZ & CARCAVALLO 1977 VenezuelaA. malheiroi SERRA, ATZINGEN & SERRA 1980 BrazilBelminus costaricencis HERRER, LENT & WYGODZINSKY 1954 Costa Rica, Mexico B. herreri LENT & WYGODZINSKY 1979 Colombia, PanamaB. laportei LENT, JURBERG & CARCAVALLO 1995 BrazilB. peruvianus HERRER, LENT & WYGODZINSKY 1954 PeruB. pittieri OSUNA & AYALA 1993 VenezuelaB. rugulosus STÅL 1859 Colombia, VenezuelaBolbodera scabrosa VALDÉS 1910 CubaMicrotriatoma borbai LENT & WYGODZINSKY 1979 BrazilM. trinidadensis (LENT 1951) Brazil, Bolivia, Colombia,

Peru,Trinidad, VenezuelaParabelminus carioca LENT 1943 BrazilP. yurupucu LENT & WYGODZINSKY 1979 BrazilCavernicola lenti BARRETT & ARIAS 1985 BrazilC. pilosa BARBER 1937 Brazil, Colombia, Panama, Peru,

VenezuelaLinshcosteus carnifex DISTANT 1904 IndiaL. chota LENT & WYGODZINSKY 1979 IndiaL. confumus GHAURI 1976 IndiaL. costalis GHAURI 1976 IndiaL. kali LENT & WYGODZINSKY 1979 IndiaL. karupus GALVÃO, PATTERSON, ROCHA & JURBERG 2002 IndiaPsammolestes arthuri (PINTO1926) Colombia, VenezuelaP. coreodes BERGROTH 1911 Argentina, Bolivia, Brazil, ParaguayP. tertius LENT & JURBERG 1965 Brazil Rhodnius amazonicus ALMEIDA, SANTOS & SPOSINA 1973 Brazil, French GuyanaRhodnius brethesi MATTA 1919 Brazil, Colombia, VenezuelaR. colombiensis MEJIA, GALVÃO & JURBERG 1999 ColombiaR. dalessandroi CARCAVALLO & BARRETO 1976 Colombia

R. domesticus NEIVA & PINTO 1923 Brazil

R. ecuadoriensis LENT & LEÓN 1958 Ecuador, Peru

R. milesi CARCAVALLO, ROCHA, GALVÃO & JURBERG 2001 Brazil

R. nasutus STÅL 1859 Brazil

R. neglectus LENT 1954 Brazil

R. neivai LENT 1953 Colombia, Venezuela

R. pallescens BARBER 1932 Belize, Colombia, Costa Rica, Pana-ma

R. paraensis SHERLOCK, GUITTON & MILES 1977 Brazil

R. pictipes STÅL 1872 Belize, Brazil, Colombia, Ecuador,Guyana, French Guyana, Peru,Suriname, Trinidad, Venezuela

R. prolixus STÅL 1859 Bolivia, Brazil, Colombia, Costa Rica,El Salvador, Ecuador, Guatemala,Guyana, French Guyana, Honduras,Mexico, Nicaragua, Panama, Suri-name, Trinidad, Venezuela

R. robustus LARROUSSE 1927 Bolivia, Brazil, Colombia, Ecuador,French Guyana, Peru, Venezuela

R. stali LENT, JURBERG & GALVÃO 1993 Bolivia, BrazilDipetalogaster maxima (UHLER 1894) MexicoEratyrus cuspidatus STÅL 1859 Colombia, Ecuador, Guatemala,

Mexico, Panama, Peru, VenezuelaE. mucronatus STÅL 1859 Bolivia, Brazil, Colombia, Ecuador,

Guatemala, Guyana, French Guyana,Panama, Peru, Suriname, Trinidad,Venezuela

Tab. 2: Geographical distribution of triatomine species (updated from GALVÃO et al. 2003).


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Behaviour and biological aspects

Egg laying and life cycle. In general theeggs of the majority of triatomine species aredeposited free in the environment, althoughsome species have an adhesive substancewhich makes the eggs stick to the substrate(Fig. 14, 15). Triatoma infestans, a domicili-ated species that originally inhabited rodentnests, is still found in the Bolivian altiplanoand lays its eggs loose in the sites it inhabits.Triatoma platensis and Triatoma delpontei, or-nithophilous species, stick their eggs to thesubstrate of the bird nests in which theylive. Species of Rhodnius also stick their eggsto the substrate. Rhodnius prolixus, which in-habits the nest of Mycteria americana, a mi-gratory bird, can have its eggs spread whenthey are adhered to the bird’s feathers.Psammolestes arthuri oviposits in clusters,like members of such reduviid subfamilies asHarpactorinae and Apiomerinae (LENT &WYGODZINSKY 1979).

Oviposition normally occurs 10 to 30days after copulation, which can be repeat-ed several times during the adult’s lifespan,although a female fecundated just once re-mains permanently fertilized. Copulationcan last 15 to 30 minutes, and the male fer-tilizes the female by depositing sper-matophores in the female vagina; theseburst and release the spermatozoa which mi-grate to the spermathecae, where they re-main protected while awaiting the passageof the successive ova to fertilize them. Thenumber of eggs varies by species and princi-pally as a function the female’s degree offeeding. Recently laid eggs are whitish orpearly in color and change color over timeuntil they become pinkish or brownish, nearthe day of hatching. During this phase, withthe eggs transparent, one can see the blackocular spots, as the 1st stage nymphs emergefrom the egg leaving the exuviae appearing(proto-nymph).

The longevity of nymphs and adultsvaries by species and ambient conditions.Laboratory experiments generally use whatare considered ideal conditions for mostspecies, with a mean temperature close to 28°C and relative humidity around 70 %,blood feeding on adequate sources for eachspecies, and a photoperiod of 12 hours. Ofcourse in nature the insect is influenced by

Species and author CountriesHermanlentia matsunoi (FERNÁNDEZ-LOAYZA 1989) PeruMeccus bassolsae (AGUILAR, TORRES, JIMENEZ, JURBERG, GALVÃO & CARCAVALLO 1999) MexicoM. longipennis (USINGER 1939) Mexico M. mazzottii (USINGER 1941) Mexico M. pallidipennis (STÅL 1872) MexicoM. phyllosomus (BURMEISTER 1835) Mexico M. picturatus (USINGER 1939) MexicoMepraia gajardoi FRIAS, HENRY & GONZALEZ 1998 ChileM. spinolai (PORTER 1934) ChileNesotriatoma bruneri USINGER 1944 CubaN. flavida (NEIVA 1911) CubaN. obscura MALDONADO & FARR 1962 JamaicaParatriatoma hirsuta BARBER 1938 Mexico, USAPanstrongylus chinai (DEL PONTE 1929) Ecuador, Peru, VenezuelaP. diasi PINTO & LENT 1946 Bolivia, BrazilP. geniculatus (LATREILLE 1811) Argentina, Bolivia, Brazil, Colombia, Costa Ri-

ca, Ecuador, Guatemala, Guyana, French Guy-ana, Mexico, Nicaragua, Panama, Paraguay,Peru, Suriname, Uruguay, Trinidad, Venezuela

P. guentheri BERG 1879 Argentina, Bolivia, Paraguay, UruguayP. howardi (NEIVA 1911) EcuadorP. humeralis (USINGER 1939) PanamaP. lenti GALVÃO & PALMA 1968 BrazilP. lignarius (WALKER 1873) Brazil, Peru, Guyana, Suriname, VenezuelaP. lutzi (NEIVA & PINTO 1923) BrazilP. megistus (BURMEISTER 1835) Argentina, Bolivia, Brazil, Paraguay, UruguayP. rufotuberculatus (CHAMPION 1899) Argentina, Bolivia, Brazil, Colombia, Costa Ri-

ca, Ecuador, Mexico, Panama, Peru, VenezuelaP. sherlocki JURBERG, CARCAVALLO & LENT 2001 BrazilP. tupynambai LENT 1942 Brazil, UruguayTriatoma amicitiae LENT 1951 Sri LankaT. arthurneivai LENT & MARTINS 1940 BrazilT. baratai CARCAVALLO & JURBERG 2000 BrazilT. barberi USINGER 1939 MexicoT. bolivari CARCAVALLO, MARTÍNEZ & PELAEZ 1987 MexicoT. bouvieri LARROUSSE 1924 Nicobar Islands, Philippine, VietnamT. brailovskyi MARTÍNEZ, CARCAVALLO & PELAEZ 1984 MexicoT. brasiliensis NEIVA 1911 BrazilT. breyeri DEL PONTE 1929 ArgentinaT. carcavalloi JURBERG, ROCHA & LENT 1998 BrazilT. carrioni LARROUSSE 1926 Ecuador, PeruT. cavernicola ELSE & CHEONG 1977 MalaysiaT. circummaculata (STÅL 1859) Brazil, UruguayT. costalimai VERANO & GALVÃO 1958 BrazilT. deaneorum GALVÃO, SOUZA & LIMA 1967 BrazilT. delpontei ROMAÑA & ABALOS 1947 Argentina, Bolivia, Brazil, Uruguay, ParaguayT. dimidiata (LATREILLE 1811) Belice, Colombia, Costa Rica, El Salvador,

Ecuador, Guatemala, Honduras, Mexico,Nicaragua, Panama, Peru, Venezuela

T. dispar LENT 1950 Colombia, Costa Rica, Ecuador, PanamaT. eratyrusiformis DEL PONTE 1929 ArgentinaT. garciabesi CARCAVALLO, CICHERO, MARTÍNEZ, PROSEN & RONDEROS 1967 Argentina, BoliviaT. gerstaeckeri (STÅL 1859) USA, Mexico T. gomeznunezi MARTÍNEZ, CARCAVALLO & JURBERG 1994 MexicoT. guasayana WYGODZINSKY & ABALOS 1949 Argentina, Bolivia, Paraguay


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various factors that are difficult to control inthe laboratory. Results obtained in the labo-ratory are thus merely an approximation ofwhat occurs in nature, but knowledge of thebiological cycles and population dynamicsallows an estimation of the species’ growthand colonization capacity, principally foranthropophilic species, but also for so-calledsecondary species with a tendency towardsdomiciliation (CANALE et al. 1999).

Temperature. The importance of tem-perature and humidity for triatomine biolo-gy was studied early by NEIVA (1913) whoobserved that high temperatures acceleratedthe embryonic period of Triatoma infestans.CARCAVALLO & MARTÍNEZ (1972) obtainedshorter cycles in specimens of three speciesof Triatoma reared at high temperatures ascompared to those reared at variable tem-peratures. SILVA (1985, 1988, 1989a, 1989b,1989c, 1990a, 1990b, 1992), SILVA & SILVA

(1988a, 1988b, 1988c, 1988d, 1988e, 1989,1990a, 1990b, 1991, 1993) and SILVA et al.(1995) compared the developmental timesof several species reared at 25 °C and 30 °Cshowing a reduction of approximately 30days in three species of Rhodnius and from40 to 60 days in Triatoma species. Accordingto most authors, when associated with lowrelative humidity the cycle is shortened bymetabolic alteration and dehydration, in-creasing the number of blood meals to bal-ance the energy budget and water loss. Var-ious authors have demonstrated accelera-tion in the developmental period of severalspecies submitted to increased temperature(GALVÃO et al. 1995, 1999b; ROCHA et al.1994, 2001a, 2001b). These results supportthe hypothesis that higher temperatures andlower relative humidity, as possible conse-quences of global warming, could acceleratethe life cycle. The result is a change in thepopulation dynamics of some Chagas diseasevectors, extending the geographic distribu-tion towards more temperate regions as wellincreasing the density of some populations.The literature shows the results of dozens ofobservations on the life cycles of these in-sects in the laboratory, a summary of theprincipal studies was published by CANALE

et al. (1999).

Species and author CountriesT. guazu LENT & WYGODZINSKY 1979 BrazilT. hegneri MAZZOTTI 1940 MexicoT. incrassata USINGER 1939 USA, Mexico T. indictiva NEIVA 1912 USA, MexicoT. infestans infestans (KLUG 1834) Argentina, Bolivia, Brazil, Chile, Ecuador,

Paraguay, Peru, UruguayT. infestans melanosoma MARTÍNEZ, OLMEDO & CARCAVALLO 1987 ArgentinaT. jurbergi CARCAVALLO, GALVÃO & LENT 1998 BrazilT. klugi CARCAVALLO, JURBERG, LENT & GALVÃO 2001 BrazilT. lecticularia (STÅL 1859) USA, MexicoT. lenti SHERLOCK & SERAFIM 1967 BrazilT. leopoldi (SCHOUDETEN 1933) Australia, IndonesiaT. limai DEL PONTE 1929 ArgentinaT. maculata (ERICHSON 1848) Aruba, Brazil, Bonaire, Curacao, Colombia,

Guyana, French Guyana, Suriname, VenezuelaT. matogrossensis LEITE & BARBOSA 1953 BrazilT. melanocephala NEIVA & PINTO1923 BrazilT. mexicana (HERRICH-SCHAEFFER 1848) MexicoT. migrans BREDDIN 1903 India, Indonesia, (Borneo, Java, Sumatra),

Malaysia, Philippine, ThailandT. neotomae NEIVA 1911 USA, MexicoT. nigromaculata (STÅL 1872) Colombia, Peru, Venezuela T. nitida USINGER 1939 Costa Rica, Guatemala, Honduras, MexicoT. oliveirai (NEIVA, PINTO & LENT 1939) BrazilT. patagonica DEL PONTE 1929 Argentina, UruguayT. peninsularis USINGER 1940 MexicoT. petrochiae PINTO & BARRETO 1925 Brazil T. platensis NEIVA 1913 Argentina, Paraguay, UruguayT. protracta (UHLER 1894) USA, MexicoT. pseudomaculata CORRÊA & ESPÍNOLA 1964 BrazilT. pugasi LENT 1953 Indonesia (Java)T. recurva (STÅL 1868) USA, MexicoT. rubida (UHLER 1894) USA, MexicoT. rubrofasciata (DE GEER 1773) Andaman Islands, Angola, Argentina, Azores, Ba-

hamas, Brazil, Burma, Cambodia, Caroline Islands,China, Comores, Congo, Cuba, Dominican Repub-lic, Formosa, French Guyana, Goa, Grenada, Gua-daloupe, Haiti, Hawaii, Hong Kong, India, Indone-sia (Borneo, Java, Sumatra) Jamaica, Madagascar,Malaysia, Martinica, Mauritius Islands, NewGuinea, Okinawa, Philippine, Saint Croix, SaintVincent, Saudi Arabia, Seychelles, Serra Leone,Singapure, South Africa, Sri Lanka, Thailand,USA,Venezuela, Vietnam, Zanzibar

T. rubrovaria (BLANCHARD 1843) Argentina, Brazil, UruguayT. ryckmani ZELEDÓN & PONCE 1972 Costa Rica, Guatemala, HondurasT. sanguisuga (LECONTE 1855) USA, Mexico T. sherlocki PAPA, JURBERG, CARCAVALLO, CERQUEIRA & BARATA 2002 BrazilT. sinaloensis RYCKMAN 1962 MexicoT. sinica HSIAO 1965 ChinaT. sordida (STÅL 1859) Argentina, Bolivia, Brazil, Paraguay, UruguayT. tibiamaculata (PINTO 1926) Brazil T. vandae CARCAVALLO, JURBERG, ROCHA, GALVÃO, NOIREAU & LENT 2002 BrazilT. venosa (STÅL 1872) Bolivia, Colombia, Costa Rica, Ecuador, PeruT. vitticeps (STÅL 1859) BrazilT. williami GALVÃO, SOUZA & LIMA 1965 BrazilT. wygodzinskyi LENT 1951 Brazil


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Fig. 11: Belminus herreri LENT & WYGODZINSKY

1979, fifth instar nymph.Fig. 12: Lateral view, head of fifth instar nymph of Linshcosteus karupus GALVÃO,PATTERSON, ROCHA, & JURBERG 2002.

Fig. 13: Rural dwelling infested by triatomine bugs.

Fig. 15: Triatominae eggs: right-free eggs (Triatomabrasiliensis NEIVA 1911).

Fig. 14: Triatominae eggs: agglutinated eggs (Rhodnius brethesiMATTA 1919).


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Feeding habits. As obligatory hemato-phages in all stages of their developmentand in both sexes, triatomines require nu-merous blood meals to complete their devel-opment. The vast majority of species feedon the blood of mammals or birds, but somecan feed on reptile or amphibian blood(CARCAVALLO et al. 1998b). Coprophagy(ingestion of feces), kleptohematophagy(sucking blood already ingested by anothertriatomine), and hemolymphagy (suckinghemolymph from other arthropods) have al-so been reported by various authors (LA-FONT 1912; BRUMPT 1914; LENT & MARTINS

1940; WOOD 1941; RYCKMAN 1951; SAN-DOVAL et al. 2000). LOROSA et al. (2000)and RUAS-NETO et al. (2001) showed thathemolymphagy can be an important sur-vival strategy in nature, observing T. cir-cummaculata and T. rubrovaria sucking blat-tids (cockroaches) in natural rock piles.Laboratory experiments by these authorsdemonstrated that the biological cycle ofboth species can be completed exclusivelywith hemolymph from these insects; howev-er, after the imaginal molt, the males and fe-males of both species could not survive onhemolymph. According to some authors,this aspect could be correlated with studieson the evolution of these two species, whichstill show this ancestral predatory character-istic.

Some species are considered stenophag-ic, that is, adapted to feeding only on givenhosts, but the vast majority have eclecticfeeding habits. CARCAVALLO et al. (1998b)published an extensive list correlating eachspecies to its respective food source. Foodsources can be identified through tech-niques like precipitin (SIQUEIRA 1960; FRE-ITAS et al. 1960), but the results may reflectmuch more the predominance of a givenhost or hosts in an area than a true food“preference“. It is common to detect thepresence of blood from various hosts in asingle insect. This result can lead to differ-ent interpretations. Could it mean greatspecies mobility? host mobility? absence ofpreference? opportunism? These questionscan only be answered safely by associatingthese results with those obtained throughethological studies. Feeding behavior de-pends on various types of stimuli in order toinduce hematophagy. Various studies have

demonstrated that heat and carbon dioxideare the principal stimuli involved in thesearch for and biting of the host (BOTTO-MAHAN et al. 2002), but heat appears to actonly on the search for food and does not in-terfere in the feeding itself. The thermore-ceptors are concentrated mainly in the an-tennae, which perform characteristic move-ments in the presence of a heat source. Bi-lateral antennectomy results in the impossi-bility of locating a heat source. Accordingto LAZZARI & NÚÑEZ (1989), it is possible tomake Triatoma infestans nymphs suck coldblood, as long as the bite is induced by ther-mal stimulation of the antennae. Anothertype of observation, aimed at characterizingthe species with greatest vector potential, isto check the number of bites/meals, dura-tion of the meal, and defecation site. Theseaspects are highly relevant epidemiological-ly, because the more contacts that occur be-tween vectors and hosts, the greater theprobability of infection or transmission ofTrypanosoma cruzi (ROCHA et al. 1997).

Resistance to long periods of fasting andthe fact that many species are generalists fa-vor their survival in nature. This capacity ofthe triatomines has been known for decades.In 1926 Uribe reported the survival of a 3rd

instar nymph of Rhodnius prolixus for fivemonths. The literature shows discrepanciesin the survival periods of various speciesstudied according to the methodology em-ployed, which can show variation in thefeeding sources, relative humidity, and tem-perature, as well as the stress the insect suf-fers during handling (WOOD 1951; FRIEND

& SMITH 1977; MASCARENHAS 1990;GALVÃO et al. 2001a; DIAS-LIMA & SHER-LOCK 2002; MARTINEZ-IBARRA et al. 2003).Resistance to fasting can vary between andwithin species. Among the different stages,the 4th and 5th are normally most resistant,because of their higher capacity to ingestblood. Various authors have focused theirlaboratory studies on the resistance periodto fasting among the various species, includ-ing GALVÃO et al. (1996, 1999a) and JU-RBERG & COSTA (1989a, 1989b).

Locomotor activity. According toBROWNE (1975) there are two forms of loco-motor activity, one spontaneous (circadian),apparently without interference from exter-


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nal stimuli, and another oriented by exter-nal stimuli. Triatomines display variousprocesses with temporal modulation, andspontaneous locomotor activity is one ofthese processes that can be observed indi-vidually and can be considered a true circa-dian rhythm. Various authors used acto-graphic records to demonstrate that locomo-tor activity in both adults and nymphs is in-tensified during the early hours of the noc-turnal period (SCHOFIELD 1976, NÚÑEZ

1982, SETTEMBRINI 1984, LAZZARI 1992).According to LAZZARI (1992), the circadianrhythm of spontaneous locomotor activity isdivided into two well demarcated moments:the search for food occurs in late after-noon/dusk and the search for shelter occursat dawn. This hypothesis was tested subse-quently by LORENZO & LAZZARI (1998),who filmed the locomotor activity of insectsin an arena containing refuges and conclud-ed that Triatoma infestans demonstratedgreater activity and motivation to feed inthe early hours of the evening than at theend of this period.

Aggregation. Aggregative behavior intriatomines is mediated principally by theresponse to the presence of chemical signals(VELÁZQUEZ ANTICH 1968). Studies per-formed with Triatoma infestans and Rhodniusprolixus demonstrated the existence of gre-garious behavior as a response to volatilesubstances found in feces (SCHOFIELD &PATTERSON 1977; ONDARZA et al. 1986).Aggregation mediated by chemical sub-stances contained in feces was recentlydemonstrated by PIRES et al. (2002) inPanstrongylus megistus and by VITTA et al.(2002) in Triatoma pseudomaculata. Despitethe various attempts at analysis, the natureof the chemical signal present in feces is stillnot well known, but it is known that fecesact as signalers for the insects’ refuges.These results show the need for future re-search aimed at the characterization of aninterspecific gregarious chemical com-pound, aimed at aiding the control andmonitoring of triatomine populations.

Camouflage. The ability of Triatomadimidiata nymphs to camouflage themselveswith particles of dust from the soil was de-scribed in detail by HASE (1940). ZELEDÓN

et al. (1969) observed the same species in

both the field and the laboratory and calledattention to this behavior’s epidemiologicalimportance. According to ZELEDÓN et al.(1973), this behavior is present in varioustriatomine species, but to variable degrees,and it appears to be completely absent inothers. One can clearly establish a correla-tion between the habitats where somespecies live (in contact with as opposed todistant from the ground) and the presenceor absence of this behavior. This behavior’sepidemiological implications deserve fur-ther investigation, since the behavior inter-feres directly with the efficiency of controlmeasures, and prior knowledge of thespecies presenting this behavior is necessaryin order to develop entomological surveil-lance methodologies.

Dispersion. Dispersion can occur pas-sively, done involuntarily by humans, or ac-tively, through the adult insects’ flight(GALVÃO et al. 2001b). Knowledge is stilllimited concerning the mechanisms in-volved in dispersion by flight. The insectsapparently respond directly to external con-ditions, but not to an internal clock. This isan extremely relevant aspect, because areasthat have been chemically treated and arefree of triatomine foci can be recolonized byflying specimens. In Triatoma infestans, themean flight distance is some 200 meters(SCHOFIELD & MATTHEWS 1985), but flightsof more than 1 kilometer have been ob-served in the field (SCHWEIGMANN et al.1988). Triatomine flight capacity has beenstudied both in the field and the laboratory;nearly all these studies were based on releas-ing and recapturing the insects, and havefurnished important information on themost flight-worthy species. However, no ex-periments have been done so far focusing onobservations of flight behavior itself(LEHANE & SCHOFIELD 1981; SCHOFIELD etal. 1991, 1992; GALVÃO et al. 2001b).

Sexual behavior. The first informationon copulation in triatomines was publishedby NEIVA (1914), who noted thatPanstrongylus megistus females appeared tocopulate only once, maintaining the eggsfertile throughout their own lifespan. Court-ing is not complex in this group of insects,and copulation in some species has been ob-served in the laboratory by some authors.


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Prior to copulation the male approaches thefemale, attempting to immobilize her withthe three legs on one side of the body in adorsolateral position (ABALOS &WYGODZINSKY 1951; HACK & BAR 1979;LENT & WYGODZINSKY 1979; LIMA et al.1986; ROJAS et al. 1990; MANRIQUE & LAZ-ZARI 1994). The presence of sexualpheromones in triatomines has been the tar-get of research for many years, and evidenceof chemical attraction between males andfemales was found in Rhodnius prolixus byVELAZQUEZ ANTICH (1968) and in Triatomainfestans and Panstrongylus megistus byNEVES & PAULINI (1981). According toBALDWIN et al. (1971), sexual pheromonesare released during copulation, leading to anaggregation of males around the couple.Similar conclusions were obtained by MAN-RIQUE & LAZZARI (1994), studying Triatomainfestans. On the other hand, the completeabsence of sexual attraction was demon-

strated by SCHOFIELD & MOREMAN (1979),HACK & BAR (1979), and LIMA & MAC-CORD (1994). Differing methodologiesprobably explain what are often contrastingresults.

Hybridization. An interesting aspect isthe hybridization capacity of these insectsunder natural conditions. Many species livein sympatry in nature and can produce in-terfertile progenies up to F2, while those thatdo not extend beyond F1 are partially inter-fertile. Ecological barriers act as an impor-tant obstacle against the meeting of differ-ent populations that are superimposed in ge-ographic areas. However, this possibility hasled various researchers to conduct hybridiza-tions in the laboratory. Table 3 shows the re-sults obtained by various authors in the lab-oratory (MAZZOTTI & OSÓRIO 1941; MAZ-ZOTTI 1943; USINGER 1944; ABALOS 1948;RYCKMAN 1962; USINGER et al. 1966; ES-PÍNOLA 1971; ZÁRATE & ZÁRATE 1985;GALÍNDEZ-GIRÓN et al. 1994; CARCAVALLO

et al. 1998a).

Laboratory rearing

Important information on triatomine bi-ology has been obtained through rearingand observation in the laboratory, it is notdifficult to establish and maintain coloniesof these insects. Some appropriate require-ments include the control of air temperatureand relative humidity. In the “LaboratórioNacional e Internacional de Referência emTaxonomia de Triatomíneos“ (National andInternational Reference Laboratory for Tri-atomine Taxonomy) at the Oswaldo CruzInstitute in Rio de Janeiro, the colonies aremaintained in glass crystallizers (20 cm highby 20 cm in diameter), covered with a re-duced-mesh nylon screen to avoid the es-cape of 1st instar nymphs and the entry ofpredators (microhymenoptera, ants, and spi-ders). Placed inside the crystallizers arestrips of filter paper that help absorb the hu-midity and increase the circulating area. Awooden stand placed inside each crystallizerserves as a support for the hosts (pigeons andmice), which are anesthetized and immobi-lized before being offered as the food source.Black paper strips are placed outside eachcrystallizer to limit the amount of light strik-ing the recipient.

Inter-crossing interfertile progeniesT. hegneri x T. dimidiata unfertile T. picturata x T. pallidipennis (F2)T. mazzottii x T. picturata (F2)T. phyllosoma x T. picturata (F2)T. phyllosoma x T. pallidipennis (F2)T. mazzotti x T. longipennis (F2)T. platensis x T. delpontei (F2)T. platensis x T. delpontei (F2)T. infestans x T. platensis (F2)T. infestans x T. rubrovaria (F1)T. sinaloensis x T. peninsularis (not)T. protracta x T. sinaloensis (F1)T. barberi x T. protracta (F1) T. barberi x T. rubida (F1: nymph V)T. maculata x T. sordida (F1)T. maculata x T. sordida (F1)T. maculata x T. infestans (F1) T. maculata x T. infestans (not) T. maculata x T. brasiliensis (F1)T. maculata x T. brasiliensis (not)T. maculata x T. pseudomaculata (F1, partial)T. pseudomaculata x T. sordida (F1)T. pseudomaculata x T. infestans (F1)T. petrocchiae x T. brasiliensis (one nymph)R. prolixus x R. neglectus (F1)R. prolixus x R. neglectus (not)R. prolixus x R. robustus (F2)R. pictipes x R. prolixus (Nymph III) (unpublished)T. matogrossensis x T. vandae (unpublished)

Tab. 3: Summarized information on experimental interspecificcrossing in Triatominae (based on CARCAVALLO et al. 1998a).


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Success in maintaining the colonies de-pends on adherence to the above-men-tioned items, taking into consideration thatoffering the same food source repeatedly forlong periods of time leads to deficiencies inthe insect’s development. When cleaningthe recipients, the strips of filter paper,which become soiled with the insects’ feces,are changed so the environment does notbecome overloaded, but some strips impreg-nated with feces are left in the environmentto allow recently hatched nymphs to havecontact with their natural digestive tractsymbionts, which aid in digestion of theblood.

Epidemiological importance andimplications for human health

The main importance of the subfamilyTriatominae is its capacity to transmit Try-panosoma cruzi, the causative agent of Cha-gas disease or American trypanosomiasis.All species of triatomine bugs must be re-garded as potential vectors of T. cruzi, whichinfect a wide variety of sylvatic and domes-tic mammals, but thus far only a few tri-atomine species have become well-adaptedto living in human dwellings, thereby ac-quiring epidemiological importance for hu-mans. The most important vector speciesare Triatoma infestans, T. brasiliensis, T.dimidiata, Panstrongylus megistus, and Rhod-nius prolixus. Heavy domestic infestations oftriatomines can be highly stressful, becausethese populations can reach several hundredindividuals of different stages, representing ahigh daily blood loss.

According to the World Health Organi-zation, Chagas disease is the third most im-portant parasitic disease (next to malariaand schistosomiasis), based on the resultingdisability and work limitations. In LatinAmerica it is the fourth most important dis-ease, following respiratory diseases, diarrhea,and AIDS (SCHOFIELD 1994). According tosero-epidemiological estimates, there areten to twenty million infected individuals inLatin America, and ninety million peoplelive in risk areas. Each year some five hun-dred thousand people are infected with thedisease, for which there is still no cure. Theexisting drugs are only partially effective inthe acute phase, which mainly attacks theheart and digestive tract. Some 10 % of in-

fected individuals develop clinical signs andsymptoms of chronic Chagas disease.

Occurrence of the classic form of thedisease in a given area depends on three ba-sic factors: presence of T. cruzi (the etiolog-ical agent), domiciliated triatomine bugs(vectors), and humans and other mammals(hosts) inhabiting the domiciliary environ-ment. In addition to the classic infectionmodel through contaminated triatomine fe-ces, accounting for 80 % of the infections,other mechanisms contribute to cases ofChagas disease, such as transfusion of con-taminated blood and blood products (16 %),congenital transmission from infectedmothers (2 %), and the rest from organtransplants, infection by the oral routethrough ingestion of contaminated food,and laboratory accidents (SCHOFIELD 1994).

Control

After the successful campaign to controldomestic triatomine populations in someSouth America countries, the new target ofstudies should be the species invading con-trolled areas. In recent years has been in-creased reporting of sylvatic species invad-ing human dwellings and peridomiciliaryenvironment in South American countries(COURA et al. 1999; VALENTE et al. 1998;VALENTE 1999; ALMEIDA et al. 2000; SAN-DOVAL et al. 2000, 2004; GALVÃO et al.2001b; VIVAS et al. 2001; WOLFF & CASTIL-LO 2002; SOUSA et al. 2004; SOUSA &GALVÃO 2004). The majority of these findsare adult insects; and, as mentioned previ-ously, triatomine flight represents an impor-tant form of dispersion to previously con-trolled areas. Studies on triatomine flightcapacity can also facilitate early identifica-tion of species with the tendency to invadedwellings and to allow the application of ad-equate vector surveillance. Effective surveil-lance is obviously important to avoid re-in-festation after control, or resurgence of anyvector population. Chagas disease controlshould be based on various independent butcomplementary work fronts, and has be-come a public health priority in the affectedcountries, due to epidemiological relevanceand the high financial costs for the econo-my.


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Use of insecticides. Vector control usesresidual-action insecticides, which shouldbe applied to both the inside and outsideparts of houses and outbuildings. One prob-lem is that the insecticides do not affect eggslaid in inaccessible places like cracks andcrevices in buildings. The vector speciesthat are subject to control should be consid-ered, since the biological cycle should bethe basis for systematizing the intervals andnumber of insecticide applications over thecourse of the year.

Housing improvement. This should bea primary goal, because typical poor ruraldwellings are made of mud and wattle, withthatched palm roofs, packed earthen floors,and have domestic and wild animals livingin the same environment, thus facilitatingmassive infestations of triatomines. Anti-triatomine measures include construction ofsimple housing with measures like smoothwalls without cracks, ceramic tile roofs pro-tected below with ceilings, and well organ-ized furniture and utensils to avoid the for-mation of refuges for insects.

Health education. Despite extensive re-search on Chagas disease, little informationhas been generated to teach rural communi-ties about the disease. The most effectivemeasure is a health education project target-ing rural communities and health profes-sionals. A population that is well-informedabout the disease is better prepared to pre-vent the entry and persistence of vectors,and to notify health authorities about exist-ing problems. Information about the diseaseand vectors and the means to control themshould be provided to the entire population,in the school system, on radio and televi-sion, and in community centers throughleaflets, posters, and films. Unfortunatelythis is still a distant reality in Latin Ameri-can countries.

When Carlos Chagas discovered themedical importance of triatomine bugs in1909 only 33 species were known, and it wasup to Arthur Neiva (through the OswaldoCruz Institute) to launch taxonomical stud-ies on this group by creating the embryo forthe Triatomines Laboratory that has beenoperating non-stop for 96 years. In 1989 theTriatomines Laboratory was transformed in-to the current “Laboratório Nacional e In-

ternacional de Referência em Taxonomia deTriatomíneos“ (National and InternationalReference Laboratory for Triatomine Taxon-omy) (JURBERG 1999), housing the world’slargest collection of triatomines, with some24,000 dry specimens, consisting of the Her-man Lent Collection with 9,000 specimensand the Rodolfo Carcavallo Collection with15,000 specimens. The Laboratory has alarge-scale insectary and is currently rearing43 species in some 150 crystallizers, in addi-tion to a collection kept in alcohol, with 45species. The collections remain open andcontinue to receive material and make do-nations, and the Laboratory is open to newscientific collaboration.

Acknowledgments

This work was supported by ConselhoNacional de Desenvolvimento Científico eTecnológico (CNPq), Serviço de Vigilânciaem Saúde-Ministério da Saúde (SVS-MS),Brazil.

Zusammenfassung

Die Arten der Unterfamilie Triatominae(Heteroptera, Reduviidae) sind Vektorenvon Trypanosoma cruzi (CHAGAS 1909), demErreger der “Chagas disease“ oderAmerikanische Trypanosomiasis. Alswichtige Vektoren sind Triatominae schonseit Jahrzehnten Gegenstand der Forschungverschiedener Aspekte ihrer Systematik, Bi-ologie, Ökologie, Biogeographie und Evolu-tion. In der vorliegenden Arbeit geben dieAutoren einen Überblick zum aktuellenKenntnisstand zur Biologie, Ökologie undSystematik dieser Vektoren und diskutierenderen Bedeutung für die menschlicheGesundheit.


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Address of the Authors:

Dr. José JURBERG

Dr. Cleber GALVÃO

Laboratório Nacional e Internacional deReferência em Taxonomia de Triatomíneos

Oswaldo Cruz InstituteAv. Brasil 4365, 21040-900

Rio de Janeiro, RJBrazil

E-Mail: [email protected] [email protected] (C. Galvão)

[email protected] (J. Jurberg)


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