Artículos Científicos

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Behavioural development and abdomen inflation of gynes and newly mated queens of Melipona beecheii (Apidae, Meliponinae)
(Universidad Nacional, Costa Rica, 1999-06-28) Van Veen, Johan; Sommeijer, M; Aguilar Monge, I
The behaviour of gynes of Melipona beecheii in queen-deprived colonies was studied. The period after emergence until acceptance is characterized by agonistic behaviour of workers towards the gynes. The gynes escaped from this worker aggression by hiding in the periphery of the nest, by performing rapid turn-arounds once grabbed by a worker, and „feigning death“. Between acceptance and nup tial flight, gynes spent most of their time in pushing, hiding, and antennal contact with workers, and self-grooming or food solicitation. After the nuptial flight the queen’s be havioural repertoire shifted to less pushing and food solicita tion, to an increase in standing, tapping and antennal contact. Accepted gynes had a significantly more prolonged ab domen inflation than gynes that were eliminated. An hypothesis is presented to explain how abdomen enlargement and behavioural development influences the acceptance of gynes and the establishment of a dominance relation with workers under queenless conditions.
A scientific note on the use of stingless bees for commercial pollination in enclosures
(INRA/DIB/AGIB/EDP Sciences, 2000) Slaa, Ester Judith; Sánchez Chaves, Luis Alejandro; Sandí, Miriam; Salazar, William
Las abejas sin aguijón se consideran polinizadores muy importantes en los trópicos, y se sabe que polinizan eficazmente al menos los cultivos. Sin embargo, rara vez se utilizan para la polinización comercial. Hasta donde sabemos, solo se ha publicado un estudio que utiliza abejas sin aguijón para la polinización de cultivos. En ese estudio, Nannotrigona testaceicornis se utilizó con éxito para polinizar fresas en un invernadero en Japón. Hay varias ventajas que pueden hacer que las abejas sin aguijón sean más adecuadas para la polinización de ciertos cultivos que la abeja melífera comúnmente utilizada.
Baltimora recta: a pollen and nectar plant for honey bees during dearth periods in Costa Rica
(Bee Worl, 2001-01-01) Ortiz Mora, Alberto; Ramíerez Arias, Fernando; Sánchez, Luis; Luis, Sánchez
Baltimora recta: a pollen and nectar plant for honey bees during dearth periods in Costa Rica. In the rainy season, during the short dry period that divides it, an extensive and dominant bloom of this herb is found on the pacific slope of Costa Rica, and it is an important food resource for honey bees.
Floral biology, pollination ecology and seed production of the ornamental plant Salvia splendens Sello
(Universidad Nacional, Costa Rica., 2002) Sánchez, Luis Alejandro; Picado, A.; Sommeijer, Marinus Jan; Slaa, Ester Judith
In this paper, the floral biology and pollination ecology of Salvia splendens are described. All flower characteristics (red corolla, large tubular flowers with abundant but dilute nectar) indicate that S. splendens is adapted to hummingbird pollination. Honeybees, however, were also found to be good pollinators of this plant. Apis mellifera was equally effective in pollinating S. splendens as hand pollination and open pollination, resulting in a 300% increase in seed set compared with bagged control plants. The much smaller stingless bee Tetragonisca angustula was not an effective pollinator of these flowers since during visitation its body failed to touch the stigma. Although pollination seemed not to be the only limiting factor in seed production, external pollinators enhance seed production in S. splendens, and Apis mellifera is an effective, commercially available pollination agent of this economically important ornamental plant.
Sounds in Melipona costaricensis (Apidae: Meliponini): effect of sugar concentration and nectar source distance
(INRA/DIB-AGIB/EDP Sciences, 2002) Aguilar Monge, Ingrid; Briceño, Daniel
The principal finding presented here is that Melipona costaricensis produces sounds that vary significantly in relation to varying sugar concentrations. Several components of the sounds produced, during the bees' stay inside of the hive, were positively correlated with sugar concentration: song duration, probability of sound production, duration of the long pulses and the relationship between short / long pulsed sound. Number of pulses per song was negatively correlated with sugar concentrations. The duration of all sound pulses was significantly correlated with the distance of the food source. Duration of the short pulses and interpulses were significantly affected by the distance of the food source. Sounds may contribute to the recruitment of nestmates to increasingly rich food supplies during the day. It is clear that aspects of the sound signals are correlated with profitability of food sources.
Efficacy of Piper (Piperaceae) extracts for control of common home and garden insect pests
(Entomological Society of America, 2004) Scott, Ian; Jensen, Helen R.; Nicol, R.W.; LeSage, Laurent; Bradbury, Rod; Sánchez Vindas, Pablo; Poveda, Luis Jorge; Arnason, John; Philogéne, Bernard J.R.
Extracts from three species of the plant family Piperaceae, Piper nigrum [L.], Piper guineense [Schum & Thonn], and Piper tuberculatum [Jacq.], were tested for efficacy against insects from five orders. All three species contain isobutyl amides, plant secondary compounds that act as neurotoxins in insects. These materials are considered safe to mammals because Piper spp. were used for centuries for spice and medicinal purposes. When 24-h P. nigrum LC50 values were compared between common insect pests from eastern Canada and the northeastern United States, the most sensitive species in order of increasing lethal concentration were eastern tent caterpillar, Malacosoma americanum (F.) European pine sawfly larvae, Neodiprion sertifer (Geoffroy) spindle ermine moth larvae, Yponomeuta cagnagella [Hübner] viburnum leaf beetle larvae, Pyrrhalta viburni [Paykull] stripped cucumber beetle adults, Acalymma vittatum (F.) Colorado potato beetle adults, Leptinotarsa decemlineata (Say) Japanese beetle adults, Popillia japonica [Newman] hairy chinch bug, Blissus leucopterus hirtis [Montandon]. The life stage tested was the point at which each species causes the greatest amount of damage to the host plant and the point at which most gardeners would likely choose to treat with a conventional synthetic insecticide. Greenhouse trials revealed that the pepper formulations also had a repellent activity, thus protecting plant leaves from 1) herbivory (lily leaf beetle, Lilioceris lilii [Scopoli], adults and larvae and stripped cucumber beetle adults) and 2) oviposition [European corn borer,Ostrinia nubilalis (Hübner)]. Combinations with other botanical extracts were additive at best in toxicity and repellent trials. Nontarget toxicity to beneficial invertebrates is a possibility because the P. nigrum LC50 for beneficial ladybird beetles was 0.2%. P. nigrum extracts can provide a reasonable level of control against lepidopteran and European pine sawfly larvae and also will work as a short-term repellent and feeding deterrent. It is recommended that the use of Piper extracts be restricted to small-scale spot treatments in residential areas where insect pest outbreaks have occurred.
Stingless Bees in applied pollination: practice and perspectives
(Universidad Nacional, Costa Rica, 2006-06-22) Sánchez Chaves, Luis; Hofstede, Frouke; Slaa, Ester Judith; Malagodi-Braga, Kátia
En la actualidad, el número de colonias de abejas, tanto silvestres como manejadas, está disminuyendo rápidamente, lo que provoca una preocupación mundial por los servicios de polinización. Las abejas sin aguijón desempeñan un importante papel ecológico como polinizadoras de muchas especies de plantas silvestres y parecen buenas candidatas para futuras alternativas en la polinización comercial. Este artículo examina la eficacia de las abejas sin aguijón como polinizadoras de cultivos. En los últimos seis años se ha duplicado el número de cultivos polinizados eficazmente por abejas sin aguijón, hasta alcanzar la cifra de 18 cultivos. Se ha descubierto que once especies de seis géneros de abejas sin aguijón forrajean eficazmente en condiciones cerradas, lo que indica el potencial de las abejas sin aguijón como polinizadoras de cultivos de invernadero. Se analizan las características biológicas que convierten a las abejas sin aguijón en firmes candidatas a prestar servicios comerciales de polinización, así como sus limitaciones actuales. Se examinan los efectos de la vegetación natural y de las abejas silvestres en el rendimiento de los cultivos, y se exponen argumentos de peso a favor de la conservación del hábitat.
A comparison of the reproductive ability of Varroa destructor (Mesostigmata:Varroidae) in worker and drone brood of Africanized honey bees (Apis mellifera)
(Springer Science, 2007-09-08) Calderón, Rafael A.; Zamora, Luis G.; Van Veen, Johan W.; Quesada, Mariela V.
Colony infestation by the parasitic mite, Varroa destructor is one of the most serious problems for beekeeping worldwide. In order to reproduce varroa females, enter worker or drone brood shortly before the cell is sealed. To test the hypothesis that, due to the preference of mites to invade drone brood to reproduce, a high proportion of the mite reproduction should occur in drone cells, a comparative study of mite reproductive rate in worker and drone brood of Africanized honey bees (AHB) was done for 370 mites. After determining the number, developmental stage and sex of the offspring in worker cells, the foundress female mite was immediately transferred into an uninfested drone cell. Mite fertility in single infested worker and drone brood cells was 76.5 and 79.3%, respectively. There was no difference between the groups (X 2 = 0.78, P = 0.37). However, one of the most significant differences in mite reproduction was the higher percentage of mites producing viable offspring (cells that contain one live adult male and at least one adult female mite) in drone cells (38.1%) compared to worker cells (13.8%) (X 2 = 55.4, P < 0.01). Furthermore, a high level of immature offspring occurred in worker cells and not in drone cells (X 2 = 69, P < 0.01). Although no differences were found in the percentage of non-reproducing mites, more than 74% (n = 85) of the mites that did not reproduce in worker brood, produced offspring when they were transferred to drone brood.
Virgin queen execution in the stingless bee Melipona beecheii: the sign stimulus for worker attacks
(INRA/DIB-AGIB/EDP Sciences, 2009-01-21) Jarau, Stefan; Van Veen, Johan W.; Aguilar, Ingrid; Ayasse, Manfred
Stingless bees produce considerable numbers of virgin queens over the year, most of which are superfluous and get executed by the workers. Nothing is known yet about the sign stimulus that releases the worker attacks. In the present study we investigated the queen execution process in Melipona beecheii and found in both behavioral observations and experiments with caged living virgin queens that workers are not attracted to them from a distance by means of volatile chemicals. Furthermore, worker aggression, which is obvious towards virgin queens that run through the nest excitedly with their abdomen enlarged and the wings beating, was lowered to almost zero when we made the queens “behavior-less” by experimentally killing them. Our results clearly show that the sign stimulus for releasing the execution behavior in Melipona beecheii workers is not a chemical stimulus but the virgin queens’ conspicuous behavior, which, we hypothesize, could act as a direct measure of their fitness.
Reproductive biology of Varroa destructor in Africanized honey bees (Apis mellifera)
(Experimental and applied Acaralogy Springer, 2009-10-23) Sánchez Chaves, Luis; Sommeijer, Marinus; Calderón, Rafael; Van Veen, Johan Wilhelm
Since its first contact with Apis mellifera, the population dynamics of the parasitic mite Varroa destructor varies from one region to another. In many regions of the world, apiculture has come to depend on the use of acaricides, because of the extensive damage caused by varroa to bee colonies. At present, the mite is considered to contribute to the recent decline of honey bee colonies in North America and Europe. Because in tropical climates worker brood rearing and varroa reproduction occurs all year round, it could be expected that here the impact of the parasite will be even more devastating. Yet, this has not been the case in tropical areas of South America. In Brazil, varroa was introduced more than 30 years ago and got established at low levels of infestation, without causing apparent damage to apiculture with Africanized honey bees (AHB). The tolerance of AHB to varroa is apparently attributable, at least in part, to resistance in the bees. The low fertility of this parasite in Africanized worker brood and the grooming and hygienic behavior of the bees are referred as important factors in keeping mite infestation low in the colonies. It has also been suggested that the type of mite influences the level of tolerance in a honey bee population. The Korea haplotype is predominant in unbalanced host-parasite systems, as exist in Europe, whereas in stable systems, as in Brazil, the Japan haplotype used to predominate. However, the patterns of varroa genetic variation have changed in Brazil. All recently sampled mites were of the Korea haplotype, regardless whether the mites had reproduced or not. The fertile mites on AHB in Brazil significantly increased from 56% in the 1980s to 86% in recent years. Nevertheless, despite the increased fertility, no increase in mite infestation rates in the colonies has been detected so far. A comprehensive literature review of varroa reproduction data, focusing on fertility and production of viable female mites, was conducted to provide insight into the Africanized bee host-parasite relationship.
A scientific note on virgin queen acceptance in stingless bees: evidence for the importance of queen aggression
(Springer Nature, 2010-01) Jarau, Stefan; Van Veen, Johan; Aguilar, Ingrid; Ayasse, Manfred
Stingless bees (Apidae, Meliponini), which are close relatives to honey bees (Apidae, Apini) and share with them a highly eusocial colonial organization (Michener, 2000), are remarkable for their habit of producing virgin queens throughout the year (Engels and Imperatriz-Fonseca, 1990). Most of these queens are not needed for a colony’s survival or division, however, and get executed some time after their emergence (Imperatriz-Fonseca and Zucchi, 1995).
Stingless bees (Scaptotrigona pectoralis) learn foreign trail pheromones and use them to find food
(Universidad Nacional de Costa Rica, 2010-11-04) Aguilar, Ingrid; Jarau, Stefan; Ayasse, Manfred; Reichle, Christian
Foragers of several species of stingless bees (Hymenoptera, Apidae and Meliponini) deposit pheromone marks in the vegetation to guide nestmates to new food sources. These pheromones are produced in the labial glands and are nest and species specific. Thus, an important question is how recruited foragers recognize their nestmates pheromone in the field. We tested whether naı¨ve workers learn a specific trail pheromone composition while being recruited by nestmates inside the hive in the species Scaptotrigona pectoralis. We installed artificial scent trails branching off from trails deposited by recruiting foragersand registered whether newly recruited bees follow these trails. The artificial trails were baited with trail pheromones of workers collected from foreign S. pectoralis colonies. When the same foreign trail pheromone was presented inside the experimental hives while recruitment took place a significant higher number of bees followed the artificial trails than in experiments without intranidal presentation. Our results demonstrate that recruits of S. pectoralis can learn the composition of specific trail pheromone bouquets inside the nest and subsequently follow this pheromone in the field. We, therefore, suggest that trail pheromone recognition in S. pectoralis is based on a flexible learning process rather than being a genetically fixed behaviour.
A scientific note on trail pheromone communication in a stingless bee, Scaptotrigona pectoralis (Hymenoptera, Apidae, Meliponini)
(INRA, DIB-AGIB and Springer Science, 2011-02-21) Jarau, Stefan; Hemmeter, Kathrin; Aguilar, Ingrid; Ayasse, Manfred
Foragers of eusocial stingless bees use a variety of communication mechanisms to inform their nestmates about the presence of valuable food sources and to recruit workers to the task of food collection (e.g., Barth et al. 2008). Recruitment to food sources at specific locations is particularly effective in species that use pheromones deposited by foragers at the resource and at various spots on the substrate along their flight root back towards the nest (Lindauer and Kerr 1960).
Nest-specific composition of the trail pheromone of the stingless bee Trigona corvina within populations
(Springer, 2012) John, Lena; Aguilar, Ingrid; Ayasse, Manfredo; Jarau, Stefan
Social insects have evolved highly developed communication systems, enabling them to coordinate complex interactions in their colonies. Pheromones play a major role in the coordination of many tasks. In Trigona corvina, a stingless bee that occurs in Central America, foragers use pheromones produced in their labial glands to scent mark solid substrates between a food source and their nest. Newly recruited bees subsequently follow these scent marks until they reach the food source. A recent study has revealed nestspecific differences in the composition of these trail pheromones in colonies of T. corvina, suggesting that pheromone specificity may serve to avoid competition between foragers from different nests. However, the nests used in this study came from different populations and their foragers certainly never met in the field (Jarau et al., 2010). The aim of the present study was to investigate whether differences in the trail pheromones of foragers from different nests can also be found between neighbouring colonies within populations. We analysed the composition of trail pheromones from labial gland secretions extracted from workers from nine colonies collected at three different populations in Costa Rica. The differences in pheromone composition were even more distinct between neighbouring nests within a population than between nests of different populations. This finding corroborates the hypothesis that nest specificity of trail pheromones serves to communicate the location of a food source exclusively to nestmates, thereby avoiding intraspecific competition at resources. Resource partitioning by avoiding conspecific non-nestmates is particularly adaptive for aggressive bee species, such as T. corvina.
Reproduction of Varroa destructor and offspring mortality in worker and drone brood cells of Africanized honey bees
(Springer, 2012-01-24) Calderón Fallas, Rafael A.; Ureña Rivera, Susana; Van Veen, Johan Wilhelm
Varroa destructor is known to be the most serious parasite of Apis mellifera worldwide. In order to reproduce varroa females enter worker or drone brood shortly before the cell is sealed. From March to December 2008, the reproductive rate and offspring mortality (mature and immature stages), focusing on male absence and male mortality of V. destructor, was investigated in naturally infested worker and drone brood of Africanized honey bees (AHB) in Costa Rica. Data were obtained from 388 to 403 single infested worker and drone brood cells, respectively. Mite fertility in worker and drone brood cells was 88. 9 and 93. 1%, respectively. There was no difference between the groups (X 2 = 3. 6, P = 0. 06). However, one of the most significant differences in mite reproduction was the higher percentage of mites producing viable offspring in drone cells (64. 8%) compared to worker cells (37. 6%) (X 2 = 57. 2, P < 0. 05). A greater proportion of mites in worker brood cells produced non-viable female offspring. Mite offspring mortality in both worker and drone cells was high in the protonymph stage (mobile and immobile). A significant finding was the high rate of male mortality. The worker and drone brood revealed that 23. 9 and 6. 9%, respectively, of the adult male offspring was found dead. If the absence (missing) of the male and adult male mortality are taken together the percentage of cells increased to 40. 0 and 21. 3% in worker and drone cells, respectively (X 2 = 28. 8, P < 0. 05). The absence of the male or male mortality in a considerable number of worker cells naturally infested with varroa is the major factor in our study which reduces the production of viable daughters in AHB colonies in Costa Rica.
Learnt information in species-specific ‘trail pheromone’ communication in stingless bees
(Elsevier, 2013) Reichle, Christian; Aguilar Monge, Ingrid; Ayasse, Manfred; Twele, Robert; Francke, Wittko; Jarau, Stefan
Specificity in biological signalling systems is often important to keep information private. Foragers of several species of stingless bees deposit chemical marking signals to guide nestmates to food sources. The markings show species- and colony-specific compositions and primarily attract a bee’s nestmates. An interesting question is whether the bees innately recognize specific trail markings or learn their particular composition from nestmates. To investigate this question, we tested whether Scaptotrigona pectoralis and Scaptotrigona subobscuripennis workers taken from their mother colonies and workers that emerged from combs transferred to foster colonies of the congeneric species are attracted to the marking compounds of workers from their natal colony or from the foster colony. A significant majority of workers were attracted to extracts prepared from foragers of the nest they inhabited, regardless of whether this was the original mother or the congeneric foster colony. Thus, the preference of stingless bee workers for specific food-marking scent mixtures is not innate, but is influenced by the odour they experience within their colony. Despite marked differences in the chemical composition of the scent marks in labial gland secretions of the two investigated species they also shared some main components. We hypothesize that recruitment trail information in stingless bees is composed of one or a few key pheromone compounds acting in conjunction with an additional signature mixture that is species and colony specific and must be learnt by recruited workers.
The antimicrobial activity and microbiological safety of stingless bee honeys from Costa Rica
(International Bee Research Association, 2014) Zamora Fallas, Gabriel; Beukelman, Kees; van den Berg, Bert; Arias Echandi, María Luisa; Umaña Rojas, Eduardo; Aguilar Monge, Ingrid; Sánchez Chávez, Luis Alejandro; Fallas Matamoros, Natalia; Quarles van Ufford, Linda; Gross, Norma
Stingless bee honeys from Costa Rica possess etnopharmacological value mainly as a wound dressing. The microbiological study by APHA methods reported that 83 % of the honeys analysed had microbial counts that comply with European Pharmacopoeia's acceptance criteria for microbiological quality of non-sterile substances for pharmaceutical use. All samples reported the absence of Clostridium botulinum spores by PCR. Over 90 % of Tetragonisca angustula and Melipona beecheii honeys inhibited Pseudomonas aeruginosa and Staphylococcus aureus at minimum inhibitory concentrations lower than Medihoney™. Under the conditions tested, Medihoney™ was not active against Candida albicans, whereas 53% of T. angustula honeys rendered inhibition. The melissopalynological analyses reported a homogeneous (monofloral) botanical composition for the Meliponini honeys, which emphasizes the contribution of nectar constituents to the antimicrobial activity and provides foundation to the standardization of a desired inhibitory effect. The traditional use of Costa Rican stingless bee honey as a dressing for burns and wounds reveals the application of a proficient antiseptic agent with low health associated risks.
Comparación de la habilidad reproductiva y mortalidad del ácaro Varroa destructor en celdas con cría de obrera y zángano en abejas africanizadas de Costa Rica
(Universidad Nacional, Costa Rica, 2014-10-31) Van Veen, Johan; Chacón, Luis Zamora; Quesada, Mariela; Calderón Fallas, Rafel Ángel
Se estudió la reproducción del ácaro Varroa destructor en celdas con cría de obrera y zángano. Además se evaluó la mortalidad de la progenie de V. destructor, especialmente la muerte o ausencia del macho. El estudio se realizó de marzo a diciembre de 2010, utilizando 15 colmenas de abejas africanizadas ubicadas en Barreal de Heredia y Ciudad Colón, San José. Se examinaron 388 celdas de obrera y 403 celdas de zángano infestadas de manera natural con un ácaro adulto. La reproducción de cada ácaro se determinó analizando los siguientes parámetros: fertilidad, fecundidad, producción de hijas viables, producción de cría inmadura, producción de únicamente hijas o únicamente machos y la ausencia de reproducción. La fertilidad de V. destructor en celdas de obrera fue de un 88.9%, con un promedio de 3.2 descendientes por ácaro. Mientras que la fertilidad del ácaro en celdas de zángano correspondió a un 93.1%, produciéndose 4.0 descendientes por ácaro. En cría de obrera, el 37.6% de los ácaros produjo hijas viables, el 14.7% hijas no viables, el 4.6% produjo cría inmadura y el 32.0% cría de un solo sexo. En celdas de zángano, el 64.8% de los ácaros produjo hijas viables, el 5.2% hijas no viables, el 1.0% cría inmadura, mientras que el 22.1% cría de un solo sexo. Además, en cría de obrera se observó una alta mortalidad en los estadios de protoninfa móvil (66.4%), protoninfa inmóvil (45.2%), deutoninfa móvil (17.6%) y el macho adulto (23.9%). En el 40.0% de las celdas de obrera con reproducción, se registró la muerte o ausencia del macho. En cría de zángano la mortalidad se registró principalmente en el estadio de protoninfa móvil (78.4%) e inmóvil (42.6%). Además, la muerte o ausencia del macho se observó en el 21.3% de las celdas de zángano. La fertilidad de V. destructor fue similar en celdas de obrera y zángano. Sin embargo, el número de hijas viables producidas por el ácaro fue mayor en la cría de zángano comparado con la cría de obrera. Por otro lado, la mortalidad o ausencia del macho fue significativamente mayor en cría de obrera. Por lo anterior, se debe indicar que el ácaro V. destructor presentó un mayor éxito reproductivo en cría de zángano, siendo está cría más apta para su reproducción.
Presence of Nosema ceranae in Africanized honey bee colonies in Costa Rica
(Universidad Nacional, Costa Rica, 2015-03-25) Sánchez Chaves, Luis; Calderón, Rafael; Yanez, Orlando; Fallas, Natalia
Nosemosis is a severe disease caused by the microsporidian Nosema sp. affecting adult honey bees (Apis mellifera) (Bailey and Ball, 1991). Nosema spores infect the epithelial layer of the ventriculus and midgut of adult bees, causing digestive disorders and shortening the life span of bees, with a resulting decrease in bee population (Ritter, 2001). So far only two microsporidian parasites have been described in honey bees: N. apis and N. ceranae. Nosema apis was the first described microsporidian in honey bees (Zander, 1909)
Presence of Deformed Wing Virus and Kashmir Bee Virus in Africanized Honey Bee Colonies in Costa Rica Infested With Varroa Destructor
(International Bee Research Association, 2015-04-01) Calderón Fallas, Rafael; Veen, Johan Van; Arce, Henry G.; Esquivel, Mario E.
Newly hatched bees with damaged wings, bees crawling in front of the hive and unusually severe mortality of adult bees were observed in Africanized bee colonies in Costa Rica. A large proportion of the adult bee population in most of these colonies was infested with Varroa destructor. Samples of adult bees were sent to the Department of Entomology at Pennsylvania State University for viral analysis. This analysis showed the presence of deformed wing virus and Kashmir bee virus. The occurrence of adult bee mortality and the detection of these viruses in Africanized colonies in Costa Rica infested with V. destructor should be considered cause for concern.

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