Folia Entomológica Mexicana ISSN: Sociedad Mexicana de Entomología, A.C. México

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1 Folia Entomológica Mexicana ISSN: Sociedad Mexicana de Entomología, A.C. México Martínez Ibarra, José Alejandro; Novelo López, Mónica Blood meals to molt, feeding time and postfeeding defecation delay of Meccus pallidipennis (Stål, 1872) (Hemiptera: Reduviidae) under laboratory conditions Folia Entomológica Mexicana, vol. 43, núm. 3, diciembre, 2004, pp Sociedad Mexicana de Entomología, A.C. Xalapa, México Available in: How to cite Complete issue More information about this article Journal's homepage in redalyc.org Scientific Information System Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Non-profit academic project, developed under the open access initiative

2 Folia Entomol. Mex., 43(3): (2004) BLOOD MEALS TO MOLT, FEEDING TIME AND POSTFEEDING DEFECATION DELAY OF MECCUS PALLIDIPENNIS (STÅL, 1872) (HEMIPTERA: REDUVIIDAE) UNDER LABORATORY CONDITIONS JOSÉ ALEJANDRO MARTÍNEZ-IBARRA AND MÓNICA NOVELO-LÓPEZ Área de Entomología Médica, Centro Universitario del Sur, Universidad de Guadalajara, Colón S/N, A. P. 20, Ciudad Guzmán, Jalisco, México. Martínez-Ibarra, J. A., and M. Novelo-López Blood meals to molt, feeding time and postfeeding defecation delay of Meccus pallidipennis (Stål, 1872) (Hemiptera: Reduviidae) under laboratory conditions. Folia Entomol. Mex., 43(3): ABSTRACT. Hatching, life cycle, blood meals to molt, mortality, fecundity, feeding time and postfeeding defecation delay for each stage of Meccus pallidipennis (Stål, 1872) were evaluated under laboratory conditions and compared to available previously published studies. Seventy-two percent of 200 eggs hatched with an average incubation period of 20.6 days. Seventy five nymphs (32.5%) completed their development to adult stadium with an average developmental time of days. Average number of blood meals per stadium was 1.4. Percent mortality for the five instars was 12.5, 10.3, 8.9, 11.6, and 17.5, respectively. Number of eggs laid per female in a 9-month period averaged ±55.7. Times per stadium to get a complete blood meals averaged 10 min and up, whereas postdefecation times averaged less than 10 min in whole the stadia. According to laboratory data, M. pallidipennis could be considered a potentially efficient vector for the transmission of Trypanosoma cruzi (Chagas) to human populations in Mexico. KEY WORDS: Meccus pallidipennis, feeding, defecation behaviors. Martínez-Ibarra, J. A. y M. Novelo-López Alimentaciones sanguíneas para mudar, tiempo de alimentación y defecación postalimentación de Meccus pallidipennis (Stål, 1872) (Hemiptera: Reduviidae) bajo condiciones de laboratorio. Folia Entomol. Mex., 43(3): RESUMEN. Se realizó un estudio bajo condiciones de laboratorio para determinar diversos parámetros de la biología de Meccus pallidipennis (Stål,1872) como: porcentaje de eclosión de huevos, duración de los estadios ninfales, número necesario de alimentaciones sanguíneas para cambiar de estadio, mortalidad, fecundidad y tiempos de alimentación y defecación postalimentación; los datos obtenidos fueron comparados con aquellos datos de estudios publicados previamente. Se obtuvo un porcentaje de eclosión de 72% en 200 huevos estudiados, con un periodo de incubación de 20.6 días. Setenta y cinco ninfas (32.5%) alcanzaron el estadio adulto, en un tiempo promedio de días. El número promedio de alimentaciones sanguíneas por estadio fue de 1.4. Los porcentajes promedio de mortalidad para los cinco estadios ninfales fueron de 12.5, 10.3, 8.9, 11.6 y 17.5, respectivamente. El número promedio de huevos puestos por hembra en un periodo de nueve meses fue de ±55.7. Los tiempos promedio de alimentación fueron de 10 min o mayores, en tanto que los tiempos promedio de defecación fueron inferiores a 10 min. Meccus pallidipennis podría ser considerado un vector potencialmente eficiente en la transmisión de Trypanosoma cruzi (Chagas) a las poblaciones humanas en México. PALABRAS CLAVE: Meccus pallidipennis, comportamiento, alimentación, defecación.

3 Martínez-Ibarra and Novelo-López: Biology of Meccus pallidipennis under laboratory conditions Chagas disease is given little public health priority in Mexico, despite ample clinical evidence of its importance (Salazar-Schettino et al.,1988; Velasco-Castrejón et al. 1991; Vidal-Acosta et al., 2000; Guzmán-Bracho, 2001; Tay et al., 2002). The etiological agent of the disease, Trypanosoma cruzi (Chagas), is transmitted directly by infected blood but most commonly by faeces of blood-sucking triatomines (Tay et al., 2002; WHO, 2002). In Mexico, at least 32 species of triatomines are known, of which 23 are unrecorded from other countries (Zárate and Zárate, 1985; WHO, 1991). Among them, Meccus (former Triatoma) pallidipennis (Stål) (see Carcavallo et al., 2000; Hypsa et al., 2002, for supporting the revalidation of the genus Meccus) is considered one of the most important vectors of Chagas disease in Mexico (Ibáñez-Bernal and Paz- Rodríguez, 1999; Vidal-Acosta et al., 2000; Guzmán-Bracho, 2001) because usually has high entomological indexes (WHO, 1991) and it occurs in houses and chicken roosts in villages of 11 states (33%) of central, southern, eastern and western Mexico (Cortés-Jiménez et al., 1996; Magallón-Gastélum et al., 1998; Bautista et al., 1999; Ramsey et al., 2000; Vidal-Acosta et al., 2000). The objective of this research was to study some unrecorded and unpublished biological parameters (v. gr. feeding time and postfeeding defecation delay) as well as to study the life cycle, mortality and the number of blood meals to molt of Meccus pallidipennis from western Mexico in order to compare it with the data from this species from central Mexico (Martínez-Ibarra and Katthain-Duchateau,1999) since some morphological differences has been noticed between populations (Martínez-Ibarra, unpublished data, Espinoza-Gómez, pers. comm.) of these two geographical areas, which could be associated to different behaviors, as reported for T. dimidiata (Latreille) (Zeledón et al., 1977, Martínez-Ibarra 2001a). As a part of a larger study on the vector efficiency of M. pallidipennis, results of studies on the vector population of M. pallidipennis under experimental conditions are presented. MATERIAL AND METHODS In order to obtain the eggs to develop our experiment, twenty individuals recently collected in Cuauhtémoc, Colima were placed in pairs, maintained at 27±1/C and 75±5% relative humidity (RH) and fed every seven days on immobilized New Zealand rabbits, since this feeding frequency has been susccesfully used to grow some related Mexican Triatominae species (Martínez-Ibarra et al. 2003a, b). Eggs were grouped by date of oviposition to initiate a cohort of 200 eggs. After eclosion, firstinstar nymphs were placed individually in plastic containers (13 cm diameter x 9 cm height), with an upcenter support of absorbent cardboard. Three days after eclosion, the nymphs were offered a blood meal individually on immobilized New Zealand rabbits during a 1-hr period until the first blood meal; subsequently, they were offered a blood meal weekly. The bugs were maintained in a dark incubator at 27±1/C and 75 ±5% RH and were checked daily for ecdysis or death. Nymphs were observed from the beginning of feeding through one hour postfeeding. Feeding time and time between blood meal and defection were recorded. From the insects that reached adult stadium, 20 couples were placed in individual containers (13 cm diameter x 9 cm height) and maintained as previously described to determine ovipositional patterns. RESULTS Seventy-two percent (n = 144/200) of the eggs hatched with an incubation period of days (20.6 ±4.1). Seventy five of these nymphs completed development to adult (Table l). Total de- 314

4 Folia Entomol. Mex., 43(3) (2004) Table 1 Development of Meccus pallidipennis fed weekly on rabbits under laboratory conditions Duration in days Stadium (n) Minimum Maximum 0± SD Egg First 144 a ± 3.1 First Second ± 4.4 Second Third ± 6.3 Third Fourth ± 11.5 Fourth Fifth ± 11.2 Fifth Adult ± 13.2 Total ± 17.3 a 200 eggs were laid Table 2 Number of blood meals and stage mortality of Meccus pallidipennis under laboratory conditions Stadium (n) Blood meals % Mortality Minimum Maximum 0 ± SD First ± Second ± Third ± Fourth ± Fifth ± Total velopmental time from egg to adult was days (204.6 ±17.3 days) (Table 1). The studied specimens took 1-6 blood meals per nymphal stadium (Table 2). Mortality was higher during the first and the fifth instar (Table 2). Mean feeding periods varied from 10 min on first and third instar to 20 min on adults (Table 3). Postfeeding defecation behavior averaged less than 10 min in all instars (Table 4). Number of eggs laid per female in a 9-month period averaged ±55.7. DISCUSSION Development of a triatomine is influenced greatly by species, environmental conditions, and the availability and kind of blood sources (Braga et al., 1998; Guarneri et al., 1998, 2000). Under laboratory conditions, development of M. pallidipennis from egg to adult was 204.6±17.3 days. This developmental period was longer than 315

5 Martínez-Ibarra and Novelo-López: Biology of Meccus pallidipennis under laboratory conditions Table 3 Mean feeding times in Meccus pallidipennis under laboratory conditions Stadium (n) Mean feeding time (minutes) Minimum Maximum 0 ± SD First ± 1.2 Second ± 1.1 Third ± 2.3 Fourth ± 1.1 Fifth ± 1.1 Adult Female ± 11.3 Adult Male ± 11.4 Table 4 Postfeeding defecation delay in Meccus pallidipennis under laboratory conditions Stadium (n) Delay (in minutes) Minimum Maximum 0± SD First ± 0.1 Second ± 2.1 Third ± 3.2 Fourth ± 4.1 Fifth ± 5.7 Adult Female ± 6.3 Adult Male ± 9.1 Rhodnius prolixus Stal (112.1 days) (Pippin, 1970), T. rubida sonoriana Usinger (119 ±6.89 days) (Paredes-González et al., in press) T. dimidiata (Latreille) (161.7 days) (Martínez-Ibarra et al., 2001) and M. pallidipennis in a previous study (187.2 ±11.71) (Martínez-Ibarra and Katthain- Duchateau, 1999), reared under similar laboratory conditions. Development time of M. pallidipennis was similar to those reported for some small species of Mexican triatomines reared under similar laboratory conditions, such as T. gerstaeckeri (Stal) (213.9 days) (Pippin, 1970) and T. barberi Usinger (range days) (Zárate, 1983), and it was also similar to those reported for some species of similar size, such as M. longipennis Usinger (192.6 days, fed on hens) (Martínez-Ibarra et al., 2003a), M. picturata Usinger (196.8 days fed on rabbits and days fed on hens) (Martínez-Ibarra et al., 2003b) and M. mazzottii Usinger (236 days) (Malo et al., 1993), 316

6 Folia Entomol. Mex., 43(3) (2004) all of them considered as important vectors of T. cruzi to human populations in Mexico (Ibáñez- Bernal and Paz-Rodríguez, 1999; Vidal-Acosta et al., 2000; Guzmán-Bracho, 2001). Hatching percentages of eggs in Triatoma or Meccus species frequently vary. They generally are higher than 80% (Zeledón et al., 1970b; Rabinovich, 1972; Zárate, 1983; Cabello and Lizano, 2001; Martínez-Ibarra et al., 2003b; Paredes-González et al., in press) although percentages less than 80% have been recorded for most Mexican triatomines, such as M. mazzottii (58.7%) (Malo et al., 1993), M. pallidipennis (60%) fed on hens in a previous study (Martínez- Ibarra and Katthain-Duchateau, 1999), T. gerstaeckeri (65.9%) (Galaviz-Silva et al., 1991) T. dimidiata (66.7%) (Martínez-Ibarra et al., 2001) M. picturata fed on hens (78.1%) (Martínez- Ibarra et al., 2003b) and M. longipennis (76.9%) (Martínez-Ibarra et al., 2003a). The 71.3% in this study could reflect the similarity between conditions in the natural habitats of M. pallidipennis (Espinoza-Gómez et al. 2002, Martínez-Ibarra unpublished data) and those present laboratory conditions. Mortality percentages were similar to those reported for M. pallidipennis in a previous study (Martínez-Ibarra and Katthain-Duchateau, 1999), since grow conditions were similar. Apparently, both populations (from central and western Mexico) are similar on success on survivorship to reach adult instar. Mortality percentages were higher during the first and fifth instar, as has been recorded for some other Triatoma and Meccus species (Zeledón et al., 1970a; Rabinovich, 1972; Zárate, 1983; Martínez-Ibarra et al., 2001, 2003a, b; Paredes-González et al. in press). In the first instar, mortality apparently resulted from an incapacity to feed, because dead nymphs had no significant blood intestinal content. In contrast, mortality on fifth instar occurred mainly during the molting (92.3%). On average, approximately 75% of second through fourth stadia required 1.4 meals to molt to the next stage, similar to M. pallidipennis in a previous study (Martínez-Ibarra and Katthain-Duchateau, 1999), M. longipennis (Martínez-Ibarra et al.,2003a) and M. picturata from the same geographic area (Martínez-Ibarra et al.,2003b), since this three groups are shown to be very similar on some other recorded behaviors, maybe because they are closely genetically related (Martínez-Ibarra, unpublished data). Average feeding periods were shorter than 21 min on each instar, indicating that M. pallidipennis is a blood meal feeder similar to T. dimidiata and to T. infestans (Klug) (Zeledón et al., 1977), which are considered some of the best vectors of T. cruzi to man. Average postfeeding defecation behavior was variable between instars, as reported for some Triatoma and Meccus species, such as T. infestans (Zeledón et al., 1977), T. barberi, (Zárate et al., 1984), T. gerstaeckeri (Galaviz-Silva et al., 1991), T. dimidiata (Martínez-Ibarra et al., 2001), M. longipennis (Martínez-Ibarra et al., 2003a), M. picturata, (Martínez-Ibarra et al., 2003b) and T. r. sonoriana (Paredes-González et al., in press). Most instars defecated in 4 min or less after feeding, suggesting a potential high effectiveness of transmission of T. cruzi to human populations under natural conditions, following to Zeledón et al. (1977), who consider that the species with postfeeding defecation behavior under 10 min could constitute effective vectors of T. cruzi. Based on the results of this study, M. pallidipennis could be potentially consider an effective transmitter of T. cruzi to humans. More studies of related biological parameters (v. gr. survivorship) are necessary as well as some studies on actual distribution and entomological indices (WHO, 1991) in order to determine the importance of M. pallidipennis as a vector of T. cruzi to human po- 317

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