Biogeography of Triatominae (Hemiptera: Reduviidae) in Ecuador: implications for the design of control strategies

Chagas disease control strategies strongly depend on the triatomine vector species involved in Trypanosoma cruzi transmission within each area. Here we report the results of the identification of specimens belonging to various species of Triatominae captured in Ecuador (15 species from 17 provinces) and deposited in the entomological collections of the Catholic University of Ecuador (Quito), Instituto Oswaldo Cruz (Brazil), the Natural History Museum London (UK), the London School of Hygiene and Tropical Medicine (UK), the National Institute of Hygiene (Quito), and the Vozandes Hospital (Quito). A critical review of published information and new field records are presented. We analysed these data in relation to the life zones where triatomines occur (11 life zones, excluding those over 2,200 m altitude), and provide biogeographical maps for each species. These records are discussed in terms of epidemiological significance and design of control strategies. Findings relevant to the control of the main vector species are emphasised. Different lines of evidence suggest that Triatoma dimidiata is not native to Ecuador-Peru, and that synanthropic populations of Rhodnius ecuadoriensis in southern Ecuador-northern Peru might be isolated from their sylvatic conspecifics. Local eradication of T. dimidiata and these R. ecuadoriensis populations might therefore be attainable. However, the presence of a wide variety of native species indicates the necessity for a strong longitudinal surveillance system.

Entomological and ecological aspects of six sylvatic species of triatomines (Hemiptera, Reduviidae) from the collection of the National Biodiversity Institute of Costa Rica, Central America

A total of 797 specimens of wild adult triatomines, belonging to six species from the entomological collections of the Costa Rican National Biodiversity Institute, was studied from the standpoint of their relative abundance, as reflected by light traps, distribution in the country, seasonal variations and climatic and altitudinal preferences. Triatoma dimidiata was the most abundant species (32.9% of the total specimens), with a very extensive distribution in different ecological zones, being more common between 100 to 400 m above sea level mainly at the end of the dry season. T. dispar was the third in frequency (21.5%), with narrower distribution, more abundant between 600 to 800 m and scarce during the dry season. Panstrongylus geniculatus and P. rufotuberculatus, second and fourth in frequency (22.1% and 15.1%, respectively), were widely distributed on both the Pacific and Caribbean basins, the former being more common between 80 to 270 m all year round and the latter below 800 m mainly during the first semester. Eratyrus cuspidatus which represented only 4.9% of the insects, was also present on both basins mainly below 200 m with a tendency to be scarce during certain months of the year, and was found in all types of ecological zones. Finally, Rhodnius pallescens, the least abundant species (3.6%) was restricted to very humid areas below 20 m, on the north side and Caribbean basin. With the exception of R. pallescens, males were more commonly found than females. Some epidemiological implications related to the six species are discussed.

Adult and nymphs of Microtriatoma trinidadensis (Lent, 1951) (Hemiptera: Reduviidae) caught from peridomestic environment in Bolivia

This is the first report of adult and nymphs (20 nymphs of all stages and 4 adults) of Microtriatoma trinidadensis (Lent 1951) (Hemiptera: Reduviidae: Triatominae) collected in peridomestic environment, in the department of La Paz, Bolivia. These specimens were associated to Rhodnius stali Lent, Jurberg & Galvão 1993. The exceptional finding of M. trinidadensis in peridomestic environment, illustrates the general tendency of triatominae to adapt to human dwellings and dependences.

Trapping Triatominae in Silvatic Habitats

Large-scale trials of a trapping system designed to collect silvatic Triatominae are reported. Live-baited adhesive traps were tested in various ecosystems and different triatomine habitats (arboreal and terrestrial). The trials were always successful, with a rate of positive habitats generally over 20% and reaching 48.4% for palm trees of the Amazon basin. Eleven species of Triatominae belonging to the three genera of public health importance (Triatoma, Rhodnius and Panstrongylus) were captured. This trapping system provides an effective way to detect the presence of triatomines in terrestrial and arboreal silvatic habitats and represents a promising tool for ecological studies. Various lines of research are contemplated to improve the performance of this trapping system.

Fipronil Insecticide: Novel Application against Triatomine Insect Vectors of Chagas Disease

We investigated the efficacy and the residual effect of fipronil® against two species of triatomine bugs, Triatoma infestans and Rhodnius neglectus, in laboratory conditions measuring concentration-response and residual activity on different surfaces (dried mud and lime coated mud). Lethal concentrations (LC50,90) were determined on filter paper. The higher insecticide efficacy against R. neglectus when compared to T. infestans may be partially attributed to the differences in their biological cycles and genetic structures. Comparison with lambdacyhalothrin wettable powder showed that fipronil mortality rates (above 50%) were observed on mud blocks and lime-coated mud blocks up to 3 months when fipronil was sprayed at 100 and 200 mg a.i./m². Residual effect deeply decayed after 3 months; and at 6 months post treatment mortality was not observed. In contrast, lambdacyhalothrin showed a long lasting residual effect on both surfaces up to 6 months. Also, it should be mentioned that fipronil had a slow, but lethal activity on the triatomine bugs when wettable formulations were used on porous surfaces.

The impact of Chagas disease control in Latin America: a review

Discovered in 1909, Chagas disease was progressively shown to be widespread throughout Latin America, affecting millions of rural people with a high impact on morbidity and mortality. With no vaccine or specific treatment available for large-scale public health interventions, the main control strategy relies on prevention of transmission, principally by eliminating the domestic insect vectors and control of transmission by blood transfusion. Vector control activities began in the 1940s, initially by means of housing improvement and then through insecticide spraying following successful field trials in Brazil (Bambui Research Centre), with similar results soon reproduced in São Paulo, Argentina, Venezuela and Chile. But national control programmes only began to be implemented after the 1970s, when technical questions were overcome and the scientific demonstration of the high social impact of Chagas disease was used to encourage political determination in favour of national campaigns (mainly in Brazil). Similarly, large-scale screening of infected blood donors in Latin America only began in the 1980s following the emergence of AIDS. By the end of the last century it became clear that continuous control in contiguous endemic areas could lead to the elimination of the most highly domestic vector populations - especially Triatoma infestans and Rhodnius prolixus - as well as substantial reductions of other widespread species such as T. brasiliensis, T. sordida, and T. dimidiata, leading in turn to interruption of disease transmission to rural people. The social impact of Chagas disease control can now be readily demonstrated by the disappearance of acute cases and of new infections in younger age groups, as well as progressive reductions of mortality and morbidity rates in controlled areas. In economic terms, the cost-benefit relationship between intervention (insecticide spraying, serology in blood banks) and the reduction of Chagas disease (in terms of medical and social care and improved productivity) is highly positive. Effective control of Chagas disease is now seen as an attainable goal that depends primarily on maintaining political will, so that the major constraints involve problems associated with the decentralisation of public health services and the progressive political disinterest in Chagas disease. Counterbalancing this are the political and technical cooperation strategies such as the "Southern Cone Initiative" launched in 1991. This international approach, coordinated by PAHO, has been highly successful, already reaching elimination of Chagas disease transmission in Uruguay, Chile, and large parts of Brazil and Argentina. The Southern Cone Initiative also helped to stimulate control campaigns in other countries of the region (Paraguay, Bolivia, Peru) which have also reached tangible regional successes. This model of international activity has been shown to be feasible and effective, with similar initiatives developed since 1997 in the Andean Region and in Central America. At present, Mexico and the Amazon Region remain as the next major challenges. With consolidation of operational programmes in all endemic countries, the future focus will be on epidemiological surveillance and care of those people already infected. In political terms, the control of Chagas disease in Latin America can be considered, so far, as a victory for international scientific cooperation, but will require continuing political commitment for sustained success.

Relationship between antennal sensilla pattern and habitat in six species of Triatominae

In order to determine if habitat similarity is correlated with a similarity of sensilla pattern, we analyzed six species of Triatominae present in two biogeographic regions of Brazil: the "caatinga" and the "cerrado". In broad terms Triatoma infestans (cerrado) and T. brasiliensis (caatinga) are found in human domiciles, T. sordida (cerrado) and T. pseudomaculata (caatinga) colonize peridomestic habitats, and Rhodnius neglectus (cerrado) and R. nasutus (caatinga) inhabit palm tree crowns. The number and distribution of four sensilla types (bristles, thin and thick walled trichoidea, and basiconica) were compared in these species. Sexual dimorphism of sensilla patterns was noted in T. sordida, T. brasiliensis and T. pseudomaculata. A principal component analysis showed three main groups: (i) species that live in the palms, (ii) domiciliated species and (iii) those living in the peridomestic habitat. T. infestans almost exclusively domestic, was placed at the centre of the canonical map and some individuals of other species overlapped there. These results support the idea that the patterns of antennal sensilla are sensitive indicators of adaptive process in Triatominae. We propose that those species that inhabit less stable habitats possess more types of sensilla on the pedicel, and higher number of antennal sensilla.

Geographic distribution and morphometric differentiation of Triatoma nitida usinger 1939 (Hemiptera: Reduviidae: Triatominae) in Guatemala

Triatoma nitida was found in 14 (0.4%) out of 3,726 houses located in six departments across Guatemala, which were surveyed from 1994 to 1998 by the man-hour collection method. Compared to previous information, the distribution of T. nitida in Guatemala has increased from five to nine departments; the species is present in mild climates at altitudes from 960 to 1,500 m. Fourteen percent of the intradomestic T. nitida were infected with Trypanosoma cruzi. The species was often found in conjunction with other triatomines (T. dimidiata and Rhodnius prolixus). The domestic and peridomestic presence of T. nitida in Guatemala was rare, but occasionally this species was colonizing human-made constructions. T. nitida appears to have a low importance as Chagas disease vector in Guatemala, as indicated by its scarce presence in the domestic habitats and defecation patterns. However, it clearly has potential to become a Chagas vector so we recommend an on-going study of the intradomestic presence of T. nitida following the control programs in Guatemala. Morphometric analysis of 47 T. nitida males from three localities showed quantitative differences between the populations, which indicates that geographic distance is an important factor in the structuring of T. nitida populations.

Epidemiology of Chagas disease in Jaguaruana, Ceará, Brazil. I. Presence of triatomines and index of Trypanosoma cruzi infection in four localities of a rural area

In order to assay the triatomine infestation and domiciliation in the rural area of Jaguaruana district, state of Ceará, Brazil, we studied, from November 2000 to April 2002, 4 localities comprising 158 domiciles as a whole, with an average of 4 inhabitants/house, who are dwelling in there for more than 7 years. Most houses have tile-covered roofs and the walls built with plaster-covered bricks (57%), followed by bricks without plaster (33%), and mud walls (7.5%). A total of 3082 triatomines were captured from different locations, according to the following capture plan: (a) intradomiciles: 238 Triatoma brasiliensis, 6 T. pseudomaculata, 9 Rhodnius nasutus, and 2 Panstrongylus lutzi; (b) peridomiciles (annexes): 2069 T. brasiliensis, 223 T. pseudomaculata, 121 R. nasutus, and 1 P. lutzi; (c) wild, in carnauba palms (Copernicia prunifera): 413 R. nasutus. From the captured triatomines, 1773 (57.5%) were examined. The natural index of Trypanosoma cruzi infection ranged from 10.8% to 30.2% (average of 17%), depending on the species and the location from where the triatomines were captured.

Blood and intestinal parasites of squirrels (Rodentia: Sciuridae) in Amazonian Brazil

We report the result of an examination for blood and intestinal protozoa in 12 specimens of the red squirrel Sciurus spadiceus (Rodentia: Sciuridae) from Birroque, municipality of Plácido de Castro, state of Acre, Brazil. No parasites were detected in thin, Giemsa-stained blood films of the animals, but culture of the blood of three in Difco B45 medium blood-agar slants gave rise to isolates of epimastigotes. Inoculation of one isolate into laboratory mice resulted in the appearance of Trypanosoma cruzi-like trypomastigotes in their peripheral blood, and the other two isolates gave rise to transient infections with a T. lewisi-like parasite in inoculated mice and hamsters. The failure of the latter parasite to develop in the triatomine bug Rhodnius robustus suggests that it is probably not T. rangeli. This appears to be the first record of a T. lewisi-like trypanosome in neotropical squirrels. Oocysts of an Eimeria sp., were detected in the faeces of 10 animals (83.3%). The parasite develops in the epithelial cells of the intestine, where it may cause severe damage and sometimes results in death of the animal. No oocysts were detected in bile.

Aspects of peridomiciliary ecotopes in rural areas of Northeastern Brazil associated to triatomine (Hemiptera, Reduviidae) infestation, vectors of Chagas disease

Artificial ecotopes of 121 peridomiciliary environments in four rural localities in the state of Ceará, Brazil, were studied and the type of material of the ecotopes was identified as triatomine infestation. Two thousand two hundred and four Triatoma brasiliensis Neiva, 340 Triatoma pseudomaculata Corrêa and Espínola, 121 Rhodnius nasutus Stall, and 5 Panstrongylus lutzi (Neiva and Pinto) were captured. Out of the 323 ecotopes found (X =2.0 ± 1.8 per dwelling) such as pigpens, henhouses, corrals, perches, dovecotes, piles of roofing tiles, bricks, wood, and straw 30.3% were infested by triatomines in all different developmental stages, including eggs. A substantial number of triatomines were found in perches, however the largest infestation took place in roofing materials used in the construction of goat/sheep corrals, henhouses, and pigpens, where 98% of them were captured: 1372 triatomines were located in the roofing tile covers, 285 in the straw, 187 in the perches, 77 in the coverings of roofing tiles and straw, and 13 in the straw and wood. Among all the different pile of materials, roofing tiles were the most infested (50%) followed by bricks (38.9%) and woods (36.1%). T. brasiliensis colonized mainly brick piles (chi2=16.539; p < 0.05) and roofing tiles (chi2=5,090.58; p < 0.05); T. pseudomaculata preferred wood perches (chi2= 472.39; p < 0.05) and woodpiles (chi2=126.0 p < 0.05), and R. nasutus was principally found in roofing straw (chi2=384.43; p < 0.05). These findings suggest that triatomines tend to colonize peridomiciliary ecotopes similar to their original habitats.

A micro-spreading improvement for spermatogenic chromosomes from Triatominae (Hemiptera-Reduviidae)

Cytogenetics of triatomines have been a valuable biological tool for the study of evolution, taxonomy, and epidemiology of these vectors of Trypanosoma cruzi. Here we present a single microtube protocol that combines micro-centrifugation and micro-spreading, allowing high quality cytogenetic preparations from male gonadal material of Rhodnius prolixus and Triatoma lecticularia. The amount of cellular scattering can be modulated, which can be useful if small aggregates of synchronous cells are desired. Moreover, a higher number of slides per gonad can be obtained with fully flattened clean chromosomal spreads with minimum overlaps, optimal for classical and modern molecular cytogenetic analyses.

Experimental transmission of the parasitic flagellates Trypanosoma cruzi and Trypanosoma rangeli between triatomine bugs or mice and captive neotropical bats

Trypanosoma cruzi and Trypanosoma rangeli-like trypanosomes have been found in a variety of neotropical bat species. In this study, bats (Artibeus lituratus, Carollia perspicillata, Desmodus rotundus, Glossophaga soricina, Molossus molossus, Phyllostomus hastatus) were maintained under controlled conditions, and experiments were conducted to determine how they might become infected naturally with trypanosomes. All bats were first screened for existing infections by hemoculture and the examination of blood smears, and only apparently uninfected animals were then used in the experiments. Proof was obtained that the triatomine bug Rhodnius prolixus would readily feed upon some of the bats, and two species became infected after being bitten by bugs infected with T. rangeli. Some bats also became infected by ingesting R. prolixus carrying T. cruzi, or following subcutaneous or intragastic inoculation with fecal suspensions of R. prolixus containing T. cruzi. P. hastatus became infected after ingesting mice carrying T. cruzi. All of the bats studied inhabit roosts that may be occupied by triatomine bugs and, with the exception of D. rotundus, all also feed to at least some extent upon insects. These findings provide further evidence of how bats may play significant roles in the epidemiology of T. cruzi and T. rangeli in the New World tropics.

Chagas disease in Andean countries

The Andean Countries' Initiative (ACI) for controlling Chagas disease was officially created in 1997 within the framework of the Hipolito Unanue Agreement (UNANUE) between the Ministries of Health of Colombia, Ecuador, Peru, and Venezuela. Its objective was to interrupt transmission via vector and transfusion in the region, taking into account that there are 12.5 million people at risk in the four Andean countries forming the initiative in the area and around 3 million people are infected by Trypanosoma cruzi. The progress of control activities for the vector species present in the Andean sub-region, for different reasons, has been slow and control interventions have still not been installed in all geographical areas occupied by the target species. This has been partly due to lack of knowledge about these vector populations' biological characteristics, and consequent uncertainty about which are the appropriate control measures and strategies to be implemented in the region. The main vector species present important similarities in Venezuela and Colombia and in Ecuador and Northern Peru and they can be approached in a similar way throughout the whole regions, basing approaches on and adapting them to the current strategies being developed in Venezuela during the 1960s which have been progressively adopted in the Southern Cone and Central-American region. Additional measures are needed for keeping endemic areas free from Rhodnius prolixus silvatic populations, widely spread in the Orinoco region in Colombia and Venezuela. Regarding aetiological treatment, it is worth mentioning that (with the exception of Colombia) none of the other countries forming the ACI have registered medicaments available for treating infected young people. There are no suitable follow-up programmes in the sub-region or for treating cases of congenital Chagas disease. An integral and integrated programme encompassing all the aspects including transmission by transfusion which seems to have achieved extremely encouraging results in all countries, are urgently needed.

Current situation of Chagas disease in Central America

Chagas disease in Central America is known since 1913 when the first human case was reported in El Salvador. The other Central American countries reported their first cases between 1933 and 1967. On October 1997 was launched the Central American Initiative for Chagas Disease Control (IPCA). The objectives of this sub-regional Initiative are: (1) the elimination of Rhodnius prolixus in Central America; (2) the reduction of the domiciliary infestation of Triatoma dimidiata; and (3) the elimination of the transfusion transmission of Trypanosoma cruzi. Significant advancements being close to the elimination of R. prolixus in Central America and the control of the transfusion transmission has been a transcendent achievement for the sub-region. The main challenges that the IPCA will have in the close future are: developing effective strategies for control and surveillance of T. dimidiata; and surveillance of other emerging triatominae species like R. pallescens, T. nitida, and T. ryckmani.