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06 Perez Paraponera Clavata.Pdf Rev. Biol. Trop., 47(4): 697-709, 1999 www.ucr.ac.cr Www.ots.ac.cr www.ots.duke.edu Distribución, mortalidad y asociación con plantas, de nidos de Paraponera clavata (Hymenoptera: Formicidae) en la isla de Barro Colorado, Panamá Rolando Pérez, Richard Condit y Suzanne Lao Centro de Ciencias Forestales del Trópico, Instituto de Investigaciones Tropicales Smithsonian, Apartado 2072, Balboa, Rep. de Panamá. Te!. (507) 227-6022 ex!. 2346, Fax. (507) 232-5978, Correo electrónico: [email protected] Recibido 22-V-1998. Corregido 15-VI-1999. Aceptado l6-VI-1999. Abstract: We studied the distribution, mortality and association with plants, of 308 nests of the neotropical ant Paraponera clavata, in a pennanen! 50 hectare plot in old-growth forest on Barro Colorado island between September 1993 and February 1995. Ant nests were unifonnly dispersed throughout the plot, and significantly associated wíth the high plateau and patch of young forest in the plot. The average densíty was 6.2 nests per hectare. Mortality of nests was higher with increasing number of neighbors within 20 m, compared to those sep­ arated at greater distances. The mortality was between 13.36% and 69.64% depending on the census interval, and recruitmentbetween 22.63% and 31.72%. The nests were found in 84 plant species of 34 families, pertain­ ing to four life fonns: 76 species were trees, 5 were shrubs, 2 were palms and one was a liana. We tested for association between ant nests and lree species and tree size by examining whether nests were more common in certain categories Ihan would be expected by change. Eight species of plants were positively associated with Paraponera clavata. The ant preferentially selected trees between 8 and 63.9 cm diameterat breas!heighl. Trees and small shrubs were not associated with the nests. No association was found between ant nests and trees with extra-floral nectaries. Fifty-three percent of the nests had a Phrynus gervaisii (Amblypygi: Phrynidae) cohabit­ ing ínside. These nests had lower mortality rates Ihan the resto Key wOl'ds: Paraponera clavata, nests, distríbution, mortality, recruitment, association, extra-floral nectaries. Paraponera clavata Smith, 1958 (Hyme­ néctar y pequeños artrópodos como escaraba­ noptera: Formicidae: Ponerinae) es una gigan­ jos, larvas de lepidópteros y arañas (Young y tesca (26 mm) hormiga neotropical que se en­ Hermann 1980). Muchos autores (Hermann cuentra distribuida desde Brasil (Sao Paulo) y 1975, Janzen y Carroll 1983, Breed y Bennett Paraguay hasta la costa atlántica de Nicaragua 1985, Barrett 1985), observaron a esta hormi­ (Janzen y Carroll 1983). Esta hormiga es un ga recolectando néctar de los nectarios extra­ insecto social que vive en colonias de unos florales de Pentaclethra macroloba (Legumi­ 500 a 1500 individuos (McCluskey y Brown nosae) y en las inflorescencias de Costus spp. 1972, Janzen y Carroll 1983), construye nidos (Zingiberaceae). Es por ello que estos autores subterráneos en la base de árboles, palmas y han tratado de establecer una relación de aso­ lianas (Belk et al. 1989, Holldobler y Wilson ciación no obligatoria entre P. clavata y árbo­ 1990a). P clavata es omnívora, se alimenta de les con nectarios. 698 REVISTADE BIOLOGÍA TROPICAL La distribución de los nidos de estas hor­ la (Hubbell y Foster 1986). El resto del bosque migas está influidapor actividades territoriales de la parcela es bosque viejo con más de 500 de colonias vecinas de la misma especie (Holl­ años de edad y pocas perturbaciones (Piperno dobler y Wilson 1990b, Ryti Y Case 1992). A 1990). Hay un mapa de la parcela en el cual se escala local, la competencia es un factor de re­ tienen perfectamente localizados, enumerados gulación en las poblaciones de hormigas con placas de aluminio e identificados, todos (Holldobler y Lumsden 1980, HolldQbler y los árboles, arbustos y palmas que tienen 1 cm Wilson 1990b). Las colonias de hormig¡¡.s pue­ o más de DAP (diámetro a la altura del pecho den compararse COIl organismos sésiles, por- . == 1.30 m desde el suelo) (Hubbell y Foster que una vez establecidas, permanecen fijas en 1983, 1987, Condit et al. 1995). Hasta 1990, se un espacio dado y sobreviven por mucho tiem­ conocía un total de 313 especies de plantas po (Thurber et al. 1993). (Condit et al. 1995). Se evalua la distribución, mortalidad y Metodología del muestreo: Entre sep­ asociación con plantas de nidos de P. clavata tiembre de 1993 y febrero de 1995, censamos y en una parcela de cincuenta hectáreas de la is" marcamos todos los nidos de P. clavata en la la de Barro Colorado, para responder las si­ parcela mediante cuatro censos: dos en la esta­ guientes preguntas: ¿Cómo están distribuidos ción lluviosa y dos en la estación seca. Las fe­ los nidos en la parcela? ¿Cómo son las tasas chas de los censos fueron: censo 1 (12-17 de de mortalidad y reclutamiento general? ¿Hay septiembre 1993), censo 2 (12-17 de febrero evidencias de asociación entre los nidos de la 1994), censo 3 (12-17 de septiembre 1994) y hormiga y his plantas de la parcela? ¿Sí los censo 4 (12-17 de febrero 1995). Los nidos fue­ nidos están asociados con árboles con necta­ ron censados por columnas de 20x500 m, orien­ rios extraflorales? ¿Sí existen otros organi­ tadas de sur a norte y de una hectárea de tama­ mas que pueden cohabitar en el nido con es­ ño. Dentro de cada columna buscamos los ni­ tas hormigas? dos por cuadrantes de 20x20 m sig�iendo siem­ pre el mismo patrón. Los nidos encontrados en la base de árboles, arbustos y palmas con más MATERIALESY MÉToDOS de 1 cm de DAP, se perturbaron con la ayuda de un tubo de aluminio de 1 m de largo y 2 cm de Zona de estudio: La isla de Barro Colo­ ancho para verificar si estaban ocupados por rado está localizada entre los 9° 09' N y 79° hormigas. Con la ayuda de la base de datos del 51' W. Es una colina de 1500 hectáreas que so­ Proyecto de Dinámica del Bosque buscamos la bresale 137 m del lago Gatún en el centro del identificación, la localización y el DAP de la Canal de Panamá. Croat (1978) y Leigh et al. planta en 1990. Hicimos anotaciones sobre los . (1990) describen detalladamente el clima, la árboles que presentaron contrafuertes y tenían flora y la fauna de la isla. Nuestro estudio se nectariosextraflorales. Después del primer cen­ hizo en una parcela permanente de cincuenta so consideramos muertos los· nidos visitados hectáreas, la cual es un rectángulo de 1000 m que no presentaron hormigas. de largo por 500 m de ancho localizada en la Análisis: Estudiamos los patrones de dis­ meseta central de la isla. El sitio es relativa­ persión de los nidos considerando los tamaños mente pláno, con unas 25 hectáreas de 0-3 por de cuadrantes qúe varían de 20 x 20 m hasta . ciento de pendientes, 13 hectáreas de 3-10 por 250 x 250 m usando el índice de dispersión de ciento de pendientes, y 10 hectáreas enpen­ Morisita (Morisita 1959, Stiteler y Pati11981). dientes moderadas (10-21 por ciento) hacia el La mortalidad anual la ca1culamos empleando este y el sur. También hay un pequeño pantano la fórmula de Condit et al. (1995). Para esta­ estacional aproximadamente de 2 hectáreas en blecer la relación entre mortalidad y densidad la parte central, y una hectárea de bosque jo­ de los nidos, utilizamos la regresión logística ven que bordea el extremo noreste de la parce- múltiple (Engelman 1990), empleando para PÉREZet al.: Distribución,mortalidad y asociacióncon plantas, de nidos de Paraponera clavata 699 ello las siguientes variables: número de nidos tró 308 nidos de P. clavata en una parcela de vecinos entre 0-10 m del nido, entre 10-20 m, cincuenta hectáreas de la isla de Barro Colo­ entre 20-30 m y entre 30-40m. Determinamos rado (Fig. 1). Se registró 237 nidos en el pri­ el reclutamiento anual utilizando la siguiente mer censo, 196 en el segundo, 202 en el terce­ fórmula: r=100 [In (S+R)-In(S)]/t, donde: ro y 210 en el cuarto, la densidad promedio r=reclutamiento anual en porcentaje, S=núme­ fue de 6.2 nidos por hectárea. Los resultados ro de nidos que sobreviven entre el primer y obtenidos con el índice de Morisita (Cuadro segundo censo, R=reclutas en el segundo cen­ 1) indican que los nidos están distribuidos so, t=intervalo en años. Paraanalizar la asocia­ uniformemente (Fig. 1), guardando cierta dis­ ción de los nidos con las especies de plantas de tancia uno del otro, especialmente en el caso la parcela de cincuenta hectáreas, comparamos de cuadrantes de 20 x 20 m y 25 x 25 m, don­ la abundancia total de nidos observada con la de el índice es menor de l. Parece ser que los esperada, tomando en cuenta el número tQtal nidos se asocian de manera significativa con de individuos de las especies de plantas en la el tipo de hábitat. Encontramos que existen parcela y empleando los datos más actualiza­ más nidos de los esperados en las planicies al­ dos del censo de 1990. Aquellas especies de tas y especialmente en la sección de bosque plantas con relativamente pocos individuos joven de la parcela (Cuadro 2), en compara­ (menos de 800), se agruparon y analizaron en ción con otras áreas. conjunto de acuerdo a su forma de vida. Mortalidad y reclutamiento de los ni­ dos: La mortalidad de nidos es alta, (69.64%) 60 nidos murieron en un intervalo de cinco me­ RESULTADOS ses entre el primer y segundo censo, (26.54%) 28 en un intervalo de siete meses ,entre el se­ Distribución de los nidos: Entre sep­ gundo y tercero, y (13.36%) 11 en un intervalo tiembre de 1993 y febrero de 1995, se encon- de cinco meses entre el tercero y el cuarto.Los G> Nidos de Paraponera clavata Fig.
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