Association between environmental gradient of anthropization and phenotypic plasticity in two species of triatomines.

BACKGROUND: Triatoma garciabesi and T. guasayana are considered secondary vectors of Trypanosoma cruzi and frequently invade rural houses in central Argentina. Wing and head structures determine the ability of triatomines to disperse. Environmental changes exert selective pressures on populations of both species, promoting changes in these structures that could have consequences for flight dispersal. The aim of this study was to investigate the relationship between a gradient of anthropization and phenotypic plasticity in flight-related traits. METHODS: The research was carried out in Cruz del Eje and Ischilín departments (Córdoba, Argentina) and included 423 individuals of the two species of triatomines. To measure the degree of anthropization, a thematic map was constructed using supervised classification, from which seven landscapes were selected, and nine landscape metrics were extracted and used in a hierarchical analysis. To determine the flight capacity and the invasion of dwellings at different levels of anthropization for both species, entomological indices were calculated. Digital images of the body, head and wings were used to measure linear and geometric morphometric variables related to flight dispersion. One-way ANOVA and canonical variate analysis (CVA) were used to analyze differences in size and shape between levels of anthropization. Procrustes variance of shape was calculated to analyze differences in phenotypic variation in heads and wings. RESULTS: Hierarchical analysis was used to classify the landscapes into three levels of anthropization: high, intermediate and low. The dispersal index for both species yielded similar results across the anthropization gradient. However, in less anthropized landscapes, the density index was higher for T. garciabesi. Additionally, in highly anthropized landscapes, females and males of both species exhibited reduced numbers. Regarding phenotypic changes, the size of body, head and wings of T. garciabesi captured in the most anthropized landscapes was greater than for those captured in less anthropized landscapes. No differences in body size were observed in T. guasayana collected in the different landscapes. However, males from highly anthropized landscapes had smaller heads and wings than those captured in less anthropized landscapes. Both wing and head shapes varied between less and more anthropogenic environments in both species. CONCLUSIONS: Results of the study indicate that the flight-dispersal characteristics of T. garciabesi and T. guasayana changed in response to varying degrees of anthropization.

Relationship between genetic diversity and morpho-functional characteristics of flight-related traits in Triatoma garciabesi (Hemiptera: Reduviidae).

BACKGROUND: Triatoma garciabesi, a potential vector of the parasitic protozoan Trypanosoma cruzi, which is the causative agent of Chagas disease, is common in peridomestic and wild environments and found throughout northwestern and central Argentina, western Paraguay and the Bolivian Chaco. Genetic differentiation of a species across its range can help to understand dispersal patterns and connectivity between habitats. Dispersal by flight is considered to be the main active dispersal strategy used by triatomines. In particular, the morphological structure of the hemelytra is associated with their function. The aim of this study was to understand how genetic diversity is structured, how morphological variation of dispersal-related traits varies with genetic diversity and how the morphological characteristics of dispersal-related traits may explain the current distribution of genetic lineages in this species. METHODS: Males from 24 populations of T. garciabesi across its distribution range were examined. The cytochrome c oxidase I gene (coI) was used for genetic diversity analyses. A geometric morphometric method based on landmarks was used for morpho-functional analysis of the hemelytra. Centroid size (CS) and shape of the forewing, and contour of both parts of the forewing, the head and the pronotum were characterised. Length and area of the forewing were measured to estimate the aspect ratio. RESULTS: The morphometric and phylogenetic analysis identified two distinct lineages, namely the Eastern and Western lineages, which coincide with different ecological regions. The Eastern lineage is found exclusively in the eastern region of Argentina (Chaco and Formosa provinces), whereas the Western lineage is prevalent in the rest of the geographical range of the species. CS, shape and aspect ratio of the hemelytra differed between lineages. The stiff portion of the forewing was more developed in the Eastern lineage. The shape of both portions of the hemelytra were significantly different between lineages, and the shape of the head and pronotum differed between lineages. CONCLUSIONS: The results provide preliminary insights into the evolution and diversification of T. garciabesi. Variation in the forewing, pronotum and head is congruent with genetic divergence. Consistent with genetic divergence, morphometry variation was clustered according to lineages, with congruent variation in the size and shape of the forewing, pronotum and head.

Phenotypic variability in traits related to flight dispersal in the wing dimorphic species Triatoma guasayana.

BACKGROUND: Triatoma guasayana is considered an emerging vector of Chagas disease in the Southern Cone of South America. The presence of a triatomine population with brachypterous individuals, in which both wings are reduced, has recently been reported for this species. The aim of the present study was to determine if flight-related traits varied across populations, if these traits could explain differences in flight capacity across populations and if flight-related traits are associated with geographic and/or climatic variation. METHODS: The study involved 66 male T. guasayana specimens from 10 triatomine populations. Digital images of wing, head and pronotum were used to estimate linear and geometric morphometric variables. Variations in size and shape were analysed using one-way analysis of variance and canonical variate analysis (CVA), respectively. Mantel tests were applied to analyse the relationship between morphometric and geographic distances, and the association between size measurements was analysed using Pearson's correlation. We explored covariation between size and shape variables using partial least square analyses (PLS). The association of geographic and climatic variables with size measurements was tested using linear regression analyses. We performed PLS analyses for shape measurements. RESULTS: Wing size differed significantly across triatomine populations. The CVA showed that wing shape of the brachypterous population is well discriminated from that of the other populations. The Mantel test showed a positive and significant association between wing shape and geographic distances. The heads of the brachypterous population were significantly larger than those of the other populations. Similar to wing shape, the head shape of the brachypterous population was well discriminated from those of the other populations. Pronotum width did not show significant differences across populations. Geographic and climatic factors were associated with size and shape of both the wing and head, but not with pronotum width. CONCLUSIONS: Most of the traits related to flight dispersal varied across populations. Wing shape and head shape were found to be better markers for differentiated morphological variation across populations. Head measurements also varied in accordance with this condition. Geographic and climatic variables were associated with most of the flight-related traits.

Modelling the effect of density vegetation coverage and the occurrence of peridomestic infestation by Triatoma infestans in rural houses of northwest of Córdoba, Argentina.

To better understand the dispersion strategies of Triatoma infestans (Klug) (Hemiptera: Reduviidae, Triatominae), we evaluated the spatial effect of infested peridomicile and density vegetation cover in a historically endemic area for Chagas disease. The study was conducted in rural houses of the northwest of Córdoba province, Argentine, during 2012-2013. Active search of triatomines were made in domicile and peridomicile habitats. To characterize vegetation coverage, a thematic map was obtained considering five types of vegetation cover (closed/open forest, closed/open shrubland and cultural land). From each house we extracted the area of vegetation coverage, housing density and infested peridomiciles density. We used generalized linear models to evaluate the effect of these variables on the occurrence of infested peridomicile. According to our results, the probability of a peridomicile to be infested increases by 1.34 (95%CI [0.98; 1.90]) times more when peridomicile structures are in environments with higher housing density and by 1.25 (95%CI [0.84; 1.88]) more times when houses are surrounded by open shrublands. Among the multiple ecological determinants of peridomestic infestation, the influence of vegetation cover has been poorly studied. In this study we discussed the effect of the vegetation as a potential modulator of the dispersion strategies of T. infestans.

Factors associated with the presence of triatomines in rural areas of south Argentine Chaco.

INTRODUCTION: The domestic and peridomestic presence of Triatoma infestans depends on several factors, such as human behavior, vector behavior, ecology, and the environment. METHODS: This work was conducted in 139 domiciliary units, where triatomines were captured and risk factors in domiciles and peridomicilies were recorded. Household dwellers were interviewed to obtain information about practices regarding this disease and entomological indicators were calculated. RESULTS: Infestation indices were 59.7% for house compounds, 4.3% for domestic areas and 58.3% for the peridomestic areas. Intradomicile infestation was significantly associated with housing characteristics. The presence of chicken coops in peridomicilies was associated with an increased risk of infestation. Of the respondents, 80% did not recognize the importance of the peridomiciliary structures for triatomine control and had infested peridomicilies. CONCLUSIONS: The results show the importance of peridomiciles as refuge sites for Triatominae bugs; however, household dwellers do not perceive peridomiciles as areas that favor the presence of vectors. Actions for raising awareness about factors that favor the presence of triatomines are needed to improve the conditions of peridomiciliary environments.

Factors associated with the presence of triatomines in rural areas of south Argentine Chaco

Abstract INTRODUCTION: The domestic and peridomestic presence of Triatoma infestans depends on several factors, such as human behavior, vector behavior, ecology, and the environment. METHODS: This work was conducted in 139 domiciliary units, where triatomines were captured and risk factors in domiciles and peridomicilies were recorded. Household dwellers were interviewed to obtain information about practices regarding this disease and entomological indicators were calculated. RESULTS: Infestation indices were 59.7% for house compounds, 4.3% for domestic areas and 58.3% for the peridomestic areas. Intradomicile infestation was significantly associated with housing characteristics. The presence of chicken coops in peridomicilies was associated with an increased risk of infestation. Of the respondents, 80% did not recognize the importance of the peridomiciliary structures for triatomine control and had infested peridomicilies. CONCLUSIONS: The results show the importance of peridomiciles as refuge sites for Triatominae bugs; however, household dwellers do not perceive peridomiciles as areas that favor the presence of vectors. Actions for raising awareness about factors that favor the presence of triatomines are needed to improve the conditions of peridomiciliary environments.

Morphological and Genetic Differentiation within the Southernmost Vector of Chagas Disease: Triatoma patagonica (Hemiptera - Reduviidae).

The epidemiological importance of Chagas disease vectors largely depends on their spreading ability and adaptation to domestic habitats. Triatoma patagonica is a secondary vector of Chagas disease endemic of Argentina, and it has been found colonizing domiciles and most commonly peridomiciliary structures in several Argentine provinces and morphological variation along its distribution range have been described. To asses if population differentiation represents geographic variants or true biological species, multiple genetic and phenotypic approaches and laboratory cross-breeding were performed in T. patagonica peridomestic populations. Analyses of chromatic variation of forewings, their size and the content of C-heterochromatin on chromosomes revealed that populations are structured following a North-South latitudinal variation. Cytochrome c oxidase I mitochondrial gene (COI) nucleotide analysis showed a mean genetic distance of 5.2% between the most distant populations. The cross-breeding experiments suggest a partial reproductive isolation between some populations with 40% of couples not laying eggs and low hatching efficiency. Our findings reveal phenotypic and genetic variations that suggest an incipient differentiation processes among T. patagonica populations with a pronounced phenotypic and genetic divergence between the most distant populations. The population differentiation here reported is probably related to differential environmental conditions and it could reflect the occurrence of an incipient speciation process in T. patagonica.

Phenotypic plasticity in response to food source in Triatoma infestans (Klug, 1834) (Hemiptera, Reduviidae: Triatominae).

In the Gran Chaco region of Argentina, Bolivia, and Paraguay, vector transmission of Trypanosoma cruzi, the etiological agent of Chagas disease, is still a severe problem because, among other causes, houses are reinfested with Triatoma infestans, the main vector of T. cruzi in southern South America. A better understanding of adaptation and evolution of T. infestans populations may contribute to the selection of appropriate vector control strategies in this region. Phenotypic plasticity is essential to understand development and maintenance of morphological variation. An experimental phenotypic plasticity study was conducted to assess if blood meal source induced head shape and size variation during development in T. infestans. Eighteen full-sib families were assigned to one of two food sources (pigeon and guinea pig) to examine the effect of food source on head shape and size in all nymph instars and adults. Data were analyzed using geometric morphometric tools and phenotypic plasticity analyses. Significant differences in head shape and size were observed between adults fed on different food sources. Allometric effects at the adult stage were observed. Head size showed significant food source × family interaction for fifth-instar nymphs and adults. For head size, significant differences between food sources were observed at stages and in ontogenetic trajectory. Phenotypic plasticity expression was found for head shape and size in adults; indeed, bugs fed on guinea pigs exhibited greater changes in head shape and larger heads than those fed on pigeon. Full-sib families exhibited different patterns of phenotypic expression in response to food source. Food source × family interaction may indicate that the observed variation in phenotypic plasticity may contribute to changes in head morphometry. These results may contribute to the selection of an appropriate control strategy for T. infestans in the Gran Chaco region, since they provide evidences of morphological plasticity in this species.

Changes related to gender, geographic population and habitat in the antennal phenotype of Triatoma patagonica Del Ponte, 1929 (Hemiptera: Reduviidae).

Triatomines undergo morphological changes as an adaptive response to different habitats (wild, peridomestic, domestic, laboratory). The characterization of the antennal phenotype provides information on intraspecific variation caused by geographical origin and/or habitat. Triatoma patagonica Del Ponte, 1929 is known to occur in peridomiciles of rural areas in Argentina, where it also invades non-colonized dwellings. Here we describe and compare the antennal phenotype of T. patagonica in populations of different geographic origin, and explore possible modifications induced by laboratory rearing with the aim of investigating the range of phenotypic variation of the species for the first time. Sixty antennas of adult males and females of T. patagonica belonging to two peridomiciliary populations of different geographical origin were analyzed. Four types of sensilla were observed in three antennal segments, showing sexual dimorphism in the species. The multivariate analysis separated the populations of similar habitat (peridomicile) but different geographical origin, without showing differences between the peridomiciliary and laboratory populations of the same geographical origin. These results suggest phenotypic plasticity in T. patagonica, which would allow the species to adapt to a wide range of habitats without having a close association with a given host and its environment. The range of antennal phenotypic variation of T. patagonica would also be an indicator of its current stage of adaptation to the human environment.

Influence of the quality and quantity of blood ingested on reproductive parameters and life-span in Triatoma infestans (Klug).

In Triatominae, female fecundity and fertility may be affected by age, adult nutritional status (i.e., blood meal source and amount of blood ingested) and number of matings. Triatoma infestans (Klug) is the main vector of Chagas disease in southern South America and considering that reproductive success is intimately associated with the potential for colonizing or re-colonizing new ecotopes in endemic areas, we studied whether the blood meal source and the amount of blood ingested have influence on reproductive parameters. We constitute two groups: couples feeding regularly on guinea pigs and couples feeding regularly on pigeons. We registered quantity of blood ingested, fecundity, fertility, number of matings, days between the first feeding and mating, copula initiation, oviposition initiation and adult life-span. Results showed that females that fed on guinea pigs exhibited high fecundity and fertility, higher number of matings and they needed a lower amount of blood to form an egg. The number of matings and fecundity increased linearly and significantly with the quantity of blood ingested for both meal sources. Results from lineal regression between life-span and fecundity showed a positive and significant relation for both meal sources. The number of matings showed a positive relation with fecundity for both meal sources but significant only for guinea pigs. In T. infestans, the quantity of blood ingested could be a determinant of their reproductive efficiency. This species is mainly adapted to human dwelling and peridomestic structures where there is low host diversity. Considering that this species is in contact with mammals over other food sources, a greater reproductive success may result from an adaptation to this environment.

Feeding and defaecation behaviour of Triatoma patagonica (Del Ponte, 1929) (Hemiptera: Reduviidae).

Among the vectors of Chagas disease, Triatoma patagonica is a species in the process of adaptation to the human environment being recently registered in urban and suburban zones. However, its importance as a vector of Chagas disease is unknown. The aim of this work was to evaluate two aspects of vectorial competence: the feeding behaviour and the defaecation pattern. These processes were studied in females of T. patagonica fed ad libitum on a restrained pigeon. The results showed that the blood meal size was negatively correlated with the time of first defaecation (r = -0.42). The first defaecation was emitted before the first 10 min and defaecations during feeding were frequent. A total of 73% of females, defaecated during the first 30 min post-feeding. These results suggest that if this species subsequently colonizes the domicile, it would be capable of transmitting Trypanosoma cruzi.

Feeding and defaecation behaviour of Triatoma patagonica (Del Ponte, 1929) (Hemiptera: Reduviidae)

Among the vectors of Chagas disease, Triatoma patagonica is a species in the process of adaptation to the human environment being recently registered in urban and suburban zones. However, its importance as a vector of Chagas disease is unknown. The aim of this work was to evaluate two aspects of vectorial competence: the feeding behaviour and the defaecation pattern. These processes were studied in females of T. patagonica fed ad libitum on a restrained pigeon. The results showed that the blood meal size was negatively correlated with the time of first defaecation (r = -0.42). The first defaecation was emitted before the first 10 min and defaecations during feeding were frequent. A total of 73 percent of females, defaecated during the first 30 min post-feeding. These results suggest that if this species subsequently colonizes the domicile, it would be capable of transmitting Trypanosoma cruzi