Flight and nutritional status of sylvatic Triatoma sordida and Triatoma guasayana

Nutritional status relating to flight activity was investigated in natural populations of Triatoma sordida and T. guasayana during the dry season in the Bolivian Chaco. The peak flight activity of both species was unimodal and covered the period 61-180 min after sunset. The weight of insects was used as the indicator of nutritional status. Interspecies comparisons employing the same sex and type of capture showed a higher weight for T. guasayana. No significant difference according to weight was observed between flying insects and those collected in natural ecotopes (hollow trees and bromeliads). More than 87% of insects collected from natural ecotopes displayed flight ability under the study conditions, explaining their tendency to invade artificial structures during the dry season.     

Vectorial capacity of Triatoma guasayana (Wygodzinsky & Abalos) (Hemiptera: Reduviidae) compared with two other species of epidemic importance

Triatoma guasayana (Wygodzinsky & Abalos) is a peridomestic triatomine with epidemiological importance in Bolivia, that may play an important role in the transmission of Trypanosoma cruzi (Chagas). In this study, two parameters of vectorial capacity were evaluated: the interval of feeding-defecation time and metacyclogenesis, in adult males and females and nymphal instars II to V of T. guasayana with comparisons with Triatoma infestans (Klug) and T. sordida (Stal). The results showed a close relationship between ingestion of blood and beginning of defecation. Values were negative in T. infestans, and T. sordida for instars II, III, IV, and V and also males and females but were positive in female T. sordida. Triatoma guasayana showed only negative values for instar II. Adults and nymphs began defecation as soon as they had finished feeding and required an average of 29.8 min. The analysis of metacyclogenesis showed that T. guasayana was superior to T. infestans and T. sordida. However, the vectorial effectiveness of T. guasayana was significantly affected regarding the percentage of metacyclic trypomastigotes during instars III to V and showed a progressive increase. Females had higher proportions despite their ingestion being half that of T. infestans. The different instars of T. guasayana had a higher parasitic load than those of T. sordida and, although ingestion was 1/3 of that of female T. infestans, there was a progressive increase in metacyclic trypomastigotes in the different nymphal stages of T. guasayana that decreased in adults.     

Environmental and demographic factors determining the spatial distribution of Triatoma guasayana in peridomestic and semi-sylvatic habitats of rural northwestern Argentina

Abstract.  Triatoma guasayana (Wygodzinsky & Abalos), a sylvatic vector of Chagas' disease, occurs in natural and peridomestic habitats of the dry Chaco region of Argentina, Bolivia and Paraguay. Ten-year retrospective spatial analyses of peridomestic T. guasayana abundance in the rural community of Amamá were expanded to the neighbouring community of Trinidad in northwestern Argentina. The distribution of T. guasayana in domiciles, peridomiciles (storerooms, chicken coops and corrals) and natural habitats (bromeliads, dry cacti and logs) around houses (i.e. 'semi-sylvatic' habitats) was analysed. The distribution of the 316 T. guasayana specimens collected in domestic and peridomestic sites during 1993–2002 was significantly clustered in both communities. Searches confirmed that the spatial distribution of semi-sylvatic and peridomestic T. guasayana was determined by the joint effects of the local abundance of goats and the density of semi-sylvatic habitats. The integration of detailed entomological and demographic longitudinal data with geographic information system data, high-resolution satellite imagery, appropriate spatial and temporal analyses and field observations allowed us to infer the underlying processes determining the distribution of T. guasayana in rural communities. This approach may be applied to other sylvatic and peridomestic vectors of Chagas' disease in order to identify high-risk areas for targeted control or environmental management.     

First report of colonies of sylvatic Triatoma infestans (Hemiptera: Reduviidae) in the Paraguayan Chaco, using a trained dog

In the Gran Chaco region, control of Triatoma infestans has been limited by persistent domestic infestations despite the efforts of the Vector Control Services. In Paraguay, this region is the highest endemic area in the country, showing high levels of indoor and outdoor infestation. Although sylvatic T. infestans have been found in the Bolivian and Argentine Chaco, similar searches for sylvatic populations of this species in Paraguay had been unsuccessful over the last 20 years. Here we present a new approach to detecting sylvatic Triatominae, using a trained dog, which has successfully confirmed sylvatic populations of T. infestans and other triatomine species in Paraguay. A total of 22 specimens corresponding to dark morph forms of T. infestans were collected, and 14 were confirmed as T. infestans by the mitochondrial cytochrome B gene analysis. Through this analysis, one of which were previously reported and a second that was a new haplotype. Triatomines were captured from amongst vegetation such as dry branches and hollows trees of different species such Aspidosperma quebracho-blanco, Bulnesia sarmientoi and Stetsonia coryne. The colonies found have been small and without apparent infection with Trypanosoma cruzi. During the study, Triatoma sordida and Triatoma guasayana have also been found in ecotopes close to those of T. infestans.     

Ecological characteristics of Triatoma patagonica at the southern limit of its distribution (Chubut, Argentina)

This study aimed at identifying the ecological characteristics of Triatoma patagonica Del Ponte 1929 (Hemiptera: Reduviidae) and evaluating the epidemiological importance of this species at its southernmost distribution limit. We carried out two surveys in the Province of Chubut, in summer and in spring, 1998. In each survey, we interviewed local health care agents for triatomine recognition and reports of the bugs, followed by entomological searches in houses and peridomestic and silvatic biotopes. The presence of T. patagonica was confirmed in two of the five localities indicated by interview. In agreement with previous studies, all these positive localities were east of the 11 degrees C isotherm, within the Monte phytogeographic province. Triatomine abundance and infestation levels (in peridomiciles and peridomestic biotopes) were higher in summer than in spring, possibly reflecting adverse environmental conditions such as flood and cold winter weather. In the silvatic environment, we found three adult bugs under rocks in summer. In peridomestic sites bugs were only found associated with chickens, which were also the only blood meal source identified. Infection by Trypanosoma cruzi was not detected. We conclude that T. patagonica at its southernmost distribution limit does not represent a risk to public health due to its low abundance, lack of association with humans, and absence of T. cruzi infection.     

Characterization of Triatoma guasayana biotopes in a hardwood forest of Santiago del Estero, Argentina

Triatoma guasayana is a silvatic triatomine species distributed in Argentina, Bolivia and Paraguay. The study was performed in a secondary forest of Santiago del Estero, Argentina. The abundance of T. guasayana was evaluated by census in the following wild biotopes: quimiles (Opuntia quimilo), chaguares (dry bromeliads), logs and underground burrows. Ten biotopes of each type were dismantled in winter (August) and another 40 in summer (January); all fauna was recorded. The biotopes most infested by T. guasayana were quimiles (65%), followed by chaguares (55%), and logs (25%). Quimiles and chaguares were infested in both seasons, whereas logs were positive only in summer and burrows were never infested. Infestation and abundance were higher in summer than in winter. The biotope structure is a key factor for T. guasayana colonization. The larger number of refuges, the constant presence of blood sources and suitable inner microclimatic conditions offered by quimiles may favour the persistence of T. guasayana colonies. The richness of invertebrate fauna per type of biotope was ranked in the same order as that of T. guasayana, suggesting similar microhabitat requirements for all studied arthropods.     

Active dispersal of Triatoma infestans and other triatomines in the Argentinean arid Chaco before and after vector control interventions

Peridomestic structures are considered the main sites where Triatoma infestans (Hemiptera: Reduviidae) remain and disperse, representing the first risk factor for intradomestic invasion, even after vector control activities. This study analyzed T. infestans dispersal during vector control interventions in six rural houses of the arid Chaco (La Rioja, Argentina). Flying and walking dispersers were captured during five months of two consecutive warm seasons after insecticide spraying of intra‐ and peridomiciles. These data were compared with previous published data in the same scenario but without insecticide spraying in peridomiciles. Recorded climatic conditions were favorable for active dispersion during the study. Total number of T. infestans dispersers moving among domestic habitats decreased after insecticide spraying. Sylvatic triatomines T. guasayana, T. eratyrusiformis, T. garciabesi, and T. platensis, not targeted by insecticide spraying, were captured simultaneously within peridomestic areas and showed higher invasion pressure than T. infestans. Adult T. infestans peridomestic populations showed high nutritional status, indicating low dispersion probability. Some peridomiciles remained infested at the end of the study. However, no intradomiciles were recolonized. These results suggest that there is a low probability of intradomestic recolonization by active dispersion from peridomiciles during 15 months post‐spraying.     

Distribution and pathogenicity of Trypanosoma cruzi isolated from peridomestic populations of Triatoma infestans and Triatoma guasayana from rural Western Argentina

We assessed the distribution of Trypanosoma cruzi infection in peridomestic triatomines collected manually at a district-wide scale in rural villages around Olta, Western Argentina, and typed the isolated strains according to their pathogenicity to laboratory mice. Of 1623 triatomines examined, only 14 (0.9%) were infected with T. cruzi based on microscopical examination of feces. The prevalence of T. cruzi infection was 0.8% in Triatoma infestans, 2.3% in T. guasayana, and nil in T. garciabesi, T. platensis, and T. eratyrusiformis. Local transmission occurred in kitchens, store-rooms and goat corrals or nearby, though at very low levels. T. cruzi was detected by at least one parasitological method in 11 (79%) of 14 microscope-positive bugs. Hemoculture was the most sensitive method (67%) followed by culture of organ homogenates, histopathology or xenodiagnosis of inoculated suckling mice (55-58%), and culture of microscope-positive bug feces (46%). The evidence suggests that most of the isolated T. cruzi strains would be myotropic type III. Our study establishes for the first time that peridomestic, microscope-positive T. guasayana nymphs were actually infected with T. cruzi, and may be implicated as a putative secondary vector of T. cruzi in domestic or peridomestic sites.     

Wing geometry as a tool for studying the Lutzomyia longipalpis (Diptera: Psychodidae) complex

Toro Toro (T) and Yungas (Y) have been described as genetically well differentiated populations of the Lutzomyia longipalpis (Lutz & Neiva, 1912) complex in Bolivia. Here we use geometric morphometrics to compare samples from these populations and new populations (Bolivia and Nicaragua), representing distant geographical origins, qualitative morphological variation ("one-spot" or "two-spots" phenotypes), ecologically distinct traits (peridomestic and silvatic populations), and possibly different epidemiological roles (transmitting or nor transmitting Leishmania chagasi). The Nicaragua (N) (Somotillo) sample was "one-spot" phenotype and a possible peridomestic vector. The Bolivian sample of the Y was also "one-spot" phenotype and a demonstrated peridomestic vector of visceral leishmaniasis (VL). The three remaining samples were silvatic, "two-spots" phenotypes. Two of them (Uyuni and T) were collected in the highlands of Bolivian where VL never has been reported. The last one (Robore, R) came from the lowlands of Bolivia, where human cases of VL are sporadically reported. The decomposition of metric variation into size and shape by geometric morphometric techniques suggests the existence of two groups (N/Y/R, and U/T). Several arguments indicate that such subdivision of Lu. longipalpis could correspond to different evolutionary units.     

Geometric morphometrics and ecological niche modelling for delimitation of near-sibling triatomine species

Identifying morphologically similar triatomine species is key to Chagas' disease vector control and surveillance, but remains challenging when only qualitative phenotypic data are available. We investigated whether morphometric and ecological variation can provide additional criteria for species delimitation by combining geometric morphometrics and ecological niche modelling to characterize two near-sibling triatomine species, Triatoma sordida and Triatoma garciabesi (Reduviidae: Triatominae). We analysed size and shape variation in 231 wings and 123 heads from one T. garciabesi and three T. sordida populations. Predicted distribution maps (21 climatic variables, 324 vector occurrence points) were produced using the Maxent method. Multivariate analyses summarized morphological and ecological variation. Wings and heads of T. sordida were significantly larger and more elongated than those of T. garciabesi. Discriminant analyses separated the species, with a partial overlap between Argentinean populations. The predicted distribution of T. garciabesi included northwest Argentina (mainly arid Chaco), whereas that of T. sordida included northeast Argentina (humid Chaco) and the Brazilian Cerrado and Caatinga ecoregions. Clear ecological niche differences were observed, with T. garciabesi occupying colder and drier areas than T. sordida. Our results show how morphometric variation and niche divergence can be used to enhance operational criteria for the delimitation of phenotypically similar triatomine species.     

Seasonal variations in active dispersal of natural populations of Triatoma infestans in rural north-western Argentina

The flight dispersal of Triatoma infestans Klug (Hemiptera: Reduviidae) is one of the main mechanisms determining community re-infestation after control interventions. An empirical model of flight initiation coupled with data from a longitudinal study predicted that the flight dispersal of T. infestans would peak in summer. To test this prediction, longitudinal light trap collections were conducted during 3-8 nights in March (late summer), July (winter) and November (spring) 2003, and in March 2004 in a rural community in north-west Argentina. Following each light-trapping collection date, all peridomestic sites around light traps were inspected to assess the relative abundance and nutritional status of T. infestans at each site. A total of 21 adult and five nymph T. infestans, six Triatoma guasayana Wygodzinsky & Abalos, and nine Triatoma garciabesi Carcavallo et al. were collected in 96 light-trapping nights, whereas 696 T. infestans were collected from the peridomestic sites that surrounded the light traps. The arrival of T. infestans in the light traps occurred in 64% of catch stations and peaked in the summer surveys (10-14 bugs) compared with spring and winter surveys. When winds were < 5 km/h, the arrival of adult T. infestans at the light traps was significantly associated with maximum temperature and relative humidity. This is the first field report of seasonal variations in the flight dispersal activity of T. infestans.     

Residual effect of a micro-encapsulated formulation of organophosphates and piriproxifen on the mortality of deltamethrin resistant Triatoma infestans populations in rural houses of the Bolivian Chaco region

The Bolivian Chaco is part of the endemic region of Chagas disease and an area where pyrethroid resistant Triatoma infestans (Hemiptera: Reduviidae) populations has been reported. The World Health Organization identified these resistant populations as an important focus for research. The objective of this study was to evaluate the residual effect of a micro-encapsulated formulation containing organophosphate active ingredients and a juvenile hormone analogue (Inesfly 5A IGR) on the mortality of T. infestans. Studies took place in rural houses of the Bolivian Chaco that were treated up to 34 months before and evaluated the susceptibility to pyrethroids of the offspring of field collected insects. Thirty houses were randomly selected within three communities to carry out wall bio-assays with T. infestans nymphs. Mortality was recorded 24, 48 and 72 h after wall contact. Eggs laid by females collected in the area were used to obtain first-instar nymphs and carry out pyrethroid susceptibility tests. The wall bio-assays showed that the micro-encapsulated insecticide eliminates T. infestans populations and produces detectable mortality of insects exposed to walls treated 34 months prior to the tests. The discriminant dose of deltamethrin (0.01 mg/mL) showed 65% nymph survival, whereas at the highest tested dose (1.0 mg/mL) 14% of the nymphs survived. These results show that Inesfly 5A IGR is an appropriate tool for the elimination of intradomestic and peridomestic populations of T. infestans resistant to pyrethroids.     

Infestation of rural houses by Triatoma infestans in the region of Los Llanos (La Rioja, Argentina)

Vectorial transmission of Chagas disease has been strongly reduced in most parts of the Southern Cone countries of South America, except in the Gran Chaco region of Argentina, Bolivia, and Paraguay. Given periodical interruptions of the vector control programmes in the endemic region of the Gran Chaco of Argentina, the vectorial transmission of the disease has been increasing during the last years. From the beginning of 2004, the provincial Ministry of Health of La Rioja, Argentina, started a vector control programme to cover the rural houses of the Los Llanos area in the southwestern area of the Gran Chaco region. This article reports the result of a standardized entomological survey and insecticide application against Chagas disease vectors in the intra and peridomestic structures of the rural houses of Los Llanos. A total of 4062 houses were inspected, of which 46.8% were found to be infested by Triatoma infestans. Infestation by vector species other than T. infestans was less than 0.5%(T. eratyrusiformis and T. platensis). Intradomestic infestation was found in 27.2%, whereas peridomestic infestation was found in 39.3% of the houses. The lowest figure of intradomestic infestation was 6.6% (Department F Varela), and the highest value of intradomestic infestation was 45.1% (Department Independencia). In spite of the demonstrated success of vector control elsewhere, this study shows that the vector populations are susceptible to pyrethroid insecticides in the southern area of the Gran Chaco of Argentina, that there still are regions where rural houses show heavy infestation by T. infestans associated with big peridomestic structures and that the vectorial transmission of the Chagas disease will continue, unless a sustained and well organized vector control effort is installed in the region.     

The role of dispersal ability, climate and spatial separation in shaping biogeographical patterns of phylogenetically distant plant groups in seasonally dry Andean forests of Bolivia

Aim To assess biogeographical patterns of Acanthaceae, Bromeliaceae, Cactaceae and Pteridophyta in Bolivian Andean seasonally dry forest islands and to explain current floristic differences between these islands by means of extrinsic (precipitation, elevation) and intrinsic (dispersal ability) factors. Location Ten isolated and disjunct seasonally dry forest areas in the Bolivian Andes and the adjacent seasonal forest areas of the Chiquitanía and Chaco regions. Methods We collated species data from recently updated and revised taxonomic treatments and herbarium collections for Acanthaceae, Bromeliaceae, Cactaceae and Pteridophyta, constructed floristic distance matrices to estimate beta diversity at the study sites and subjected them to Mantel correlation analyses. Multiple regressions on distance matrices allowed us to test the influence of geographical distance and environmental (elevation and precipitation) differences on floristic differentiation. Results Acanthaceae and Bromeliaceae, and to a lesser extent Cactaceae, showed coincident biogeographical patterns and suggested the presence of two seasonally dry forest groups in Andean Bolivia: one including all small isolated northern dry valleys and another including all southern valleys with connections to the lowland seasonal forests of the Chiquitanía and the Chaco. Most of the variation in the floristic distance matrices in these plant groups, with seed dispersal typically restricted to short distances, was explained by spatial separation between habitat islands. In contrast, pteridophytes showed a different biogeographical pattern. Their floristic differences between sites were determined by the environmental variables. The anemochorous and spore‐based dispersal system of this plant group seems to be a highly effective mechanism allowing pteridophytes to easily reach even the isolated dry valleys in inter‐Andean Bolivia. Main conclusions Current biogeographical patterns in dry Andean habitat islands can provide insights into the factors that control the processes of community assembly. We show that differences in community composition of phylogenetically distant plant groups in the understorey of seasonally dry forest islands can be explained by a combination of the habitat characteristics where the group is present (either precipitation, elevation or both) and, more interestingly, by group specific dispersal limitation (as inferred by geographical distance between island habitats).     

Genetic structure of Triatoma sordida (Hemiptera: Reduviidae) domestic populations from Bolivia: application on control interventions.

The genetic population of Triatoma sordida group 1, a secondary vector of Chagas disease in Bolivia, was studied by multi-locus enzyme electrophoresis. A total of 253 nymphal and adult specimens collected from seven neighbouring localities in the Velasco Province, Department of Santa Cruz, were processed. The relatively low genetic variability was confirmed for this species (rate of polymorphism: 0.20). The absence of genetic disequilibrium detected within the seven localities was demonstrated. A geographical structuration appears between localities with distances greater than 20 km apart. Although T. sordida presents a relatively reduced dispersive capacity, its panmictic unit is wider than compared with T. infestans. Genetic distances between T. sordida populations were correlated with geographic distance. Gene flow between geographic populations of T. sordida provides an efficient framework for effective vigilance and control protocols.     

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.     

Sylvatic population of Triatoma infestans from the Bolivian Chaco: from field collection to characterization

A sylvatic Triatoma infestans DM (dark morph) population detected in the Bolivian Chaco was characterized and compared with various domestic ones. The degree of differentiation of DM was clearly within the T. infestans intra-specific level. Nevertheless marked chromatic and morphometric differences as well as differences in antennal pattern, chromosome banding and randomly amplified polymorphic DNA support the hypothesis of a distinct population. Continuous exchange of insects between wild and domestic habitats seems unlikely in the Chaco.     

Activity of oil-formulated Beauveria bassiana against Triatoma sordida in peridomestic areas in Central Brazil

Field tests were carried out during the rainy season of 2001/2002 in S?o Luís de Montes Belos, Goiás, Brazil, to evaluate the potential of the entomopathogenic fungus, Beauveria bassiana, against peridomestic Triatoma sordida. An oil-water formulation of the isolate CG 14 (Embrapa) was applied in triatomine infested hen houses of four farms at a final concentration of 10(6) conidia/cm2. Numbers of T. sordida decreased over the next 25 days, after application of the fungus, and B. bassiana developed on dead insects in one hen house. A high number of B. bassiana colonies was detected in substrates collected in treated hen houses 24 h after application of CG 14. In the following three months the presence of B. bassiana declined to values found before treatment.     

Molecular phylogeography of the Chagas' disease vector Triatoma infestans in Argentina

Triatoma infestans is the main vector of Chagas' disease in South America between latitudes 10°S and 46°S. A multilocus microsatellite data set of 836 individuals from 27 populations of T. infestans, from all its range of distribution in Argentina, was analyzed. Our results favor the hypothesis of two independent migration events of colonization in Argentina and secondary contacts. The majority of the populations of the western provinces of Catamarca, La Rioja, San Juan and the west of Cordoba province, had almost no shared ancestry with the rest of the populations analyzed. Probably those populations, belonging to localities close to the Andean region, could have been established by the dispersal line of T. infestans that would have arrived to Argentina through the Andes, whereas most of the rest of the populations analyzed may have derived from the dispersal line of T. infestans in non-Andean lowlands. Among them, those from the provinces of Formosa, Chaco, Santiago del Estero and Santa Fe shared different percentages of ancestry and presented lower degree of genetic differentiation. The migratory movement linked to regional economies and possibly associated with passive dispersal, would allow a higher genetic exchange among these populations of T. infestans. This study, using microsatellite markers, provides a new approach for evaluating the validity of the different hypotheses concerning the evolutionary history of this species. Two major lineages of T. infestans, an Andean and non-Andean, are suggested.     

Dispersal of Triatoma infestans and other Triatominae species in the arid Chaco of Argentina: flying, walking or passive carriage? The importance of walking females

The aim of this paper was to analyse the active dispersal of Triatoma infestans and the role of chickens as passive carriers of this insect in peridomestic areas of La Rioja, Argentina. To measure active dispersal, monthly catches were made on six consecutive nights for five months (in the warm season) using light traps (for flying insects) and sticky dispersal barriers (for walking insects). The nutritional and reproductive states of adults were evaluated. Over the course of the sampling period, a total of eight flying adults, six walking nymphs and 10 walking adults of the species T. infestans were captured, as well as specimens of Triatoma guasayana, Triatoma eratyrusiformis and Triatoma platensis. Our data demonstrate for the first time that females of T. infestans can disperse by walking. This may be an adaptive strategy because it allows them to move with eggs and/or with good blood reserves, which are not possible when flying. All flying and walking individuals of both genders were of an appropriate physiological state that would allow for colonisation of the target habitat. However, manual inspection of 122 chickens suggests that it is unlikely that these animals passively transport T. infestans. Finally, the dispersal activity of T. infestans was compared with other triatomines using a dispersion index.