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(Erebidae: Arctiinae)?, RCCB http://www.rccb.uh.cu arTÍCULO original ¿Es imprescindible la comunicación acústica en la conducta de apareamiento de Empyreuma pugione (Erebidae: Arctiinae)? Is accoustic communication essential in the mating behavior of Empyreuma pugione (Erebidae: Arctiinae)? Yohami Fernández,1 Martha Pérez1 y Emanuel C. Mora1* 1 Departamento de biología animal RESUMEN y Humana, Facultad de biología, La comunicación química y acústica participa en el reconocimiento y la Universidad de la Habana, Cuba. aceptación de la pareja durante la conducta de apareamiento de muchas * Autor para correspondencia: especies de lepidópteros nocturnos. En el presente trabajo, se analiza el [email protected] efecto de la audición y la emisión de sonido sobre la probabilidad de apa- rearse en Empyreuma pugione. Para ello, se combinaron hembras y machos capaces de producir y detectar sonido con individuos sordomudos y se cuantificó el número de apareamientos exitosos entre tres grupos experi- mentales: (1) hembras y machos intactos, (2) machos intactos y hembras sordomudas, (3) machos sordomudos y hembras intactas. Se observaron apareamientos exitosos en las tres combinaciones sin diferencias significa- tivas entre las tres condiciones experimentales. El 29 % de los apareamien- tos tuvo lugar entre machos sordomudos y hembras intactas, mientras el 24 % correspondió a hembras sordomudas que se aparearon con machos intactos. Este hallazgo sugiere que la emisión de sonido en E. pugione no resulta imprescindible para el éxito del apareamiento, y probablemente la comunicación mediante feromonas conjuntamente con otros sistemas sensoriales sean suficientes para seleccionar y aceptar a la pareja. PALABRAS CLAVE: comunicación acústica, apareamiento, Empyreuma pugione. ABSTRACT Chemical and acoustic communication are involved in species recognition and in female mate-choice during the mating behavior of many nocturnal Lepidoptera. Here, we analyzed the effect of audition and sound production in Empyreuma pugione reproductive behavior. For this, we combine intact females and males with deaf-mute animals and measured the number of suc- cessful matings in three experimental conditions: (1) couples of intact females and males, (2) couples of deaf-mute males and intact females, (3) couples of intact males and deaf-mute females. We found successful matings for the three tested combinations without significant difference between experimental conditions. 29 % of the successful mating occurred between deaf-mute males and intact females and 24 % between deaf-mute females and intact males. Our results suggest that sound production in E. pugione is not essential during courtship behavior and probably the chemical communication combined with other sensory systems is enough for select and accept mates. RECIBIDO: 9/2012 ACEPTADO: 11/2012 KEYWORDS: acoustic communication, mating behavior, Empyreuma pugione. REVISTA CUBANA DE CIENCIAS BIOLÓGICAS rnps: 2362 • issn: 2307-695x • vol. 2 • n.o 1 • enero-abril • 2013 • pp. 32-37 ¿es impresCinDible la ComUniCaCión aCúsTiCa en la Conducta De apareamienTo De EMPYREUMA PUGIONE (EREBIDAE: ARCTIINAE)? 33 YoHami FernánDez • martha pérez • emanUel C. mora Introducción química y la acústica participan en la conducta La liberación de feromonas desempeña un papel de apareamiento y que su eliminación reduce el esencial en el cortejo de la mayoría de las especies número de acoplamientos exitosos. Sin embargo, de lepidópteros nocturnos estudiados hasta el mo- en muchos casos, ninguna de las dos resulta im- mento. Adicionalmente, en un creciente número de prescindible para que se produzca la cópula entre especies de polillas, se ha descrito el uso de señales algunas parejas. Sobre esta base formulamos la acústicas en combinación con las feromonas para el hipótesis de que la probabilidad de apareamiento reconocimiento y la aceptación de la pareja (Nakano en E. pugione se incrementa si se favorecen los et al., 2006; Takanashi et al., 2010). La producción contactos físicos entre los individuos, independien- de sonido en este contexto ha sido demostrada temente de su capacidad de producir y detectar principalmente en las familias Pyralidae, Arctiidae, sonido. Noctuidae, Geometridae y Crambidae (Spangler En este trabajo se analiza el efecto de la audi- et al., 1984; Spangler, 1985; Snedden et al., 1994; ción y la emisión de sonido sobre la probabilidad Sanderford y Conner, 1995; Conner, 1999; Nakano de aparearse en E. pugione. Para ello, se colocaron et al., 2006, 2008, 2009a, 2009b, 2010a, 2010b; grupos de 45 ejemplares en recipientes de aproxi- Takanashi et al., 2010). De acuerdo con esos resul- madamente 25 L. En cada grupo se combinaron tados, la importancia relativa del sonido respecto a hembras y machos capaces de detectar y pro- la emisión de feromonas varía según la especie. ducir sonido, y sordomudos. Como indicador del Las características espectrales y temporales éxito en el apareamiento se valoró el número de de las señales emitidas por los órganos produc- parejas de cada tipo que se apareaban en cada tores de sonido de Empyreuma pugione (= affinis) día de observación. han sido descritas por Coro et al. (1983) y Sander- ford et al. (1998). El papel de la emisión acústica Materiales y métodos durante el apareamiento se ha investigado tanto Se utilizaron 330 individuos de la especie Empyreuma en condiciones de laboratorio como en el campo. pugione (Noctuoidea: Erebidae, Arctiinae), 247 hem- Portilla et al. (1987) analizaron el efecto de la pér- bras y 83 machos, recolectados en los municipios dida de la audición sobre el tiempo dedicado a la Playa y Plaza de la Revolución en La Habana, en actividad sexual y la probabilidad de apareamiento, estadio adulto. En el laboratorio, los ejemplares en parejas de esta especie aisladas en recipientes se mantuvieron en condiciones de temperatura, de un litro. En estas condiciones, describieron una humedad y fotoperiodo semejantes a las de su disminución del tiempo de actividad sexual y ausen- medio natural y fueron alimentados con solución cia casi total de apareamientos cuando uno de los de sacarosa al 30 % ad libitum. Los experimentos dos miembros de la pareja era sordo. Tal evidencia se llevaron a cabo entre el 12 y el 22 de diciembre de de la participación del sonido en la comunicación 2008, en el periodo de 5:30 a.m. a 8:00 a.m., que entre los sexos fue demostrada por Sanderford y es el horario del día en que esta especie se aparea Coro (1998) al registrar señales acústicas durante (Otazo et al., 1984). el cortejo de esta especie en condiciones naturales. Para comprobar la emisión de sonido durante el Estos últimos autores refieren que el macho emite apareamiento y el papel de las emisiones acústicas sonido cuando se acerca a una hembra que libera en el éxito de esta actividad, se diseñó un experi- feromona y mientras intenta copular con ella. La mento utilizando polillas intactas y sordomudas. hembra puede o no emitir sonido durante la fase La condición de sordera se logró destruyendo la táctil del cortejo. Cuando lo hace, comienza después membrana timpánica de ambos órganos auditivos de la señal del macho y se mantiene entre uno y dos con un alfiler entomológico, bajo un estereoscopio segundos después que esta cesa. ZEISS modelo KL1500 (Nakano et al., 2006, 2009a, Los resultados obtenidos en otras especies de 2010a). De forma similar, la posibilidad de emitir polillas (Conner, 1987; Sanderford y Conner, 1995; sonido se evitó rasgando la banda estriada de los Nakano et al., 2006, 2009a, 2010a, 2010b; Taka- timbales a lo largo de toda su longitud (Nakano et nashi et al., 2010) indican que la comunicación al., 2010b). Con esta manipulación, la deformación REVISTA CUBANA DE CIENCIAS BIOLÓGICAS rnps: 2362 • issn: 2307-695x • vol. 2 • n.o 1 • enero-abril • 2013 • pp. 32-37 ¿es impresCinDible la ComUniCaCión aCúsTiCa en la Conducta De apareamienTo De EMPYREUMA PUGIONE (EREBIDAE: ARCTIINAE)? 34 YoHami FernánDez • martha pérez • emanUel C. mora mecánica del episternito, que se produce al pre- individuos incapaces de establecer comunicación sionar el tórax entre los dedos índice y pulgar por acústica se comparó con la distribución de pro- la zona de inserción de las alas (Coro et al., 1983), babilidades obtenida para los individuos intactos. no originaba pulsos acústicos. Lo anterior se Se tomó como criterio de diferencia significativa comprobó con un micrófono ultrasónico colocado p < 0,05, lo cual equivale a que las probabilidades cerca de la preparación. de apareamiento fueran diferentes en más de un Una vez manipulados, los animales se separa- 95 % de las veces analizadas. ron en tres contenedores cilíndricos de malla, de 32 x 31 cm y aproximadamente 25 L de capacidad. Resultados En cada contenedor se colocaron 45 individuos Durante los once días de observación se produje- (30 hembras y 15 machos). La mayor proporción ron 34 apareamientos exitosos. De manera ge- de hembras tuvo como objetivo aumentar la neral, se aparearon tanto parejas formadas por probabilidad de que alguna de ellas emitiera individuos intactos, como parejas formadas feromonas y atrajera al macho (Nakano et al., por al menos un individuo incapaz de establecer 2009a, 2010a; Takanashi et al., 2010). El primer comunicación acústica (sordomudo). La figura 1 contenedor se conformó con individuos intactos. ilustra el número de apareamientos por día y el En el segundo se incluyeron machos intactos y total para cada tipo de pareja a lo largo de los hembras sordomudas y, en el tercero, machos once días de observación. Entre los individuos sordomudos
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