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Ecology of a Costa Rican Leaf-Cutting Ant 61 Wetterer et al.: Ecology of a Costa Rican Leaf-cutting Ant 61 FORAGING AND NESTING ECOLOGY OF ACROMYRMEX OCTOSPINOSUS (HYMENOPTERA: FORMICIDAE) IN A COSTA RICAN TROPICAL DRY FOREST JAMES K. WETTERER,1 DANIEL S. GRUNER2 AND JORGÉ E. LOPEZ3 1Center for Environmental Research & Conservation, Columbia University New York, NY 10027 2Department of Zoology, University of Hawaii, Honolulu, HI 96822 3Department of Ecology and Evolution, University of Chicago, Chicago, IL 60637 ABSTRACT Leaf-cutting ants (Acromyrmex sp. and Atta sp.) in Costa Rica show many intra- and interspecific differences in ecology. Recent taxonomic studies question whether the Acromyrmex octospinosus populations on the Pacific and Atlantic slopes of Costa Rica are a single species. We therefore examined the foraging and nesting ecology of A. octospinosus in the tropical dry forest of Palo Verde National Park on the Pacific slope of Costa Rica and compared our findings with published data on the ecology of A. octospinosus in the tropical moist forest of La Selva Biological Station on the At- lantic slope. The Pacific A. octospinosus foraged primarily on the leaves of herbs and other small plants, fallen leaves, fruit, flowers, and insect frass, but does not cut the leaves of large trees. Worker size distribution within colonies was bimodal with only 62 Florida Entomologist 81(1) March, 1998 the larger workers leaving the nest to forage. Nests were shallow and generally under a few centimeters of organic debris at the base of trees and woody shrubs or in crev- ices. The foraging and nesting ecology of the Pacific A. octospinosus appeared to be very similar to that of the Atlantic A. octospinosus, except that the Pacific ants col- lected considerable amounts of insect frass (11% of all loads), whereas the Atlantic ants had no recorded loads of frass. This difference in selectivity, however, may have been due simply to seasonal differences in availability of frass at the sites. Acromyrmex octospinosus was the only species of leaf-cutting ant found at Palo Verde. The vertisol soil of the area, which has very poor drainage when wet and cracks deeply when dry, may not be suitable for major pest species of leaf-cutters in Costa Rica, Atta cephalotes and Atta colombica, which excavate nests deep underground. Key Words: Atta, foraging selectivity, Palo Verde, pest ants RESUMEN Las hormigas podadoras (Acromyrmex sp. y Atta sp.) en Costa Rica muestran mu- chas diferencias intra- e interespecíficas en ecología. Estudios taxonómicos recientes ponen en duda la noción que poblaciones de Acromyrmex octospinosus en los flancos Orientales y Occidentales de Costa Rica sean una sola especie. Por lo tanto, nosotros examinamos la ecología de alimentaciùn y anidamiento de A. octospinosus en el bos- que seco tropical del Parque Nacional Palo Verde en el flanco occidental de Costa Rica, y comparamos nuestros resultados con datos previamente publicados sobre la ecología de A. octospinosus en el bosque húmedo tropical de la Estación Biológica La Selva en el flanco oriental. La alimentación de A. octospinosus en el lado occidental consistió principalmente de hojas de hierbas y otras plantas pequeñas, hojas caídas, frutas, flo- res y excrementos de insectos, pero no cortan hojas de árboles grandes. La distribu- ción del tamaño de las obreras en la colonia es bimodal, y sólo las obreras grandes salen de la colonia para recoger alimentos. Los nidos son poco profundos y por lo ge- neral están bajo unos cuantos centímetros de materia orgánica al pie de árboles y ar- bustos, o en grietas. La ecología de alimentación y anidamiento de A. octospinosus en el lado occidental parece ser muy similar a la de A. octospinosus en el lado oriental, ex- cepto que las hormigas en el lado occidental recogen cantidades considerables de ex- cremento de insectos (11% de todas las cargas), mientras que las hormigas en el lado oriental no han sido observadas cargando excremento de insectos. Esta diferencia en selectividad, sin embargo, puede ser causada por diferencias en disponibilidad de ex- cremento de insectos en diferentes estaciones entre las localidades. Acromyrmex oc- tospinosus fue la única especie de hormiga podadora encontrada en Palo Verde. El suelo vertisol de esa área, que tiene muy mal drenaje cuando está mojado y que forma rajaduras hondas cuando está seco, puede no ser adequado para otras especies dañi- nas de hormigas podadoras en Costa Rica, Atta cephalotes y Atta colombica, las cuales excavan nidos profundos. Fungus-growing ants (Myrmicinae; Attini) are unique among ants in their habit of cultivating a symbiotic fungus for food. All attine ants are obligately dependent on their fungus. In other aspects of their ecologies, however, different species of attines vary greatly (Weber 1972, Hölldobler & Wilson 1990). Ants of the nine “lesser” genera (Cyphomyrmex, Mycetosoritis, Mycetophylax, Mycocepurus, Mycetarotes, Myrmicoc- rypta, Apterostigma, Sericomyrmex, and Trachymyrmex) typically collect insect frass, dead insects, or small pieces of fallen plant material to use as substrate for their fun- gal gardens. They tend to have small and inconspicuous colonies (fewer than 5000 in- dividuals), with relatively monomorphic workers. In contrast, ants belonging to two Wetterer et al.: Ecology of a Costa Rican Leaf-cutting Ant 63 genera of attines, Acromyrmex and Atta, often depend on harvesting leaves for their fungal gardens and consequently are commonly called leaf-cutting ants. Leaf-cutting ant colonies can grow to include many thousand to several million highly-polymorphic workers. Many leaf-cutters, particularly Atta species, are major agricultural pests. Earlier studies have found striking differences in the foraging and nesting ecology of Acromyrmex octospinosus (Reich), Acromyrmex volcanus (Wheeler), Acromyrmex coronatus (Fabricius), and Atta cephalotes (L.), the four species of leaf-cutting ants found on the Atlantic slope of Costa Rica (Wetterer, 1991, 1993, 1994a, b, 1995). Ac- romyrmex octospinosus and A. volcanus colonies forage primarily on small herbs and fallen leaves, fruits, and flowers (Wetterer 1991, 1993). These colonies produce two distinct size castes of workers (Wetterer, unpublished). The minima workers stay in- side the nest, tending the fungus garden and brood, while the relatively monomorphic maxima workers leave the nest to forage. In A. coronatus colonies and small A. ceph- alotes colonies, workers primarily cut the soft leaves of herbaceous plants (Wetterer 1994a, 1995). These colonies produce a narrow range of small workers, the largest of which are just big enough to cut soft vegetation efficiently (Wilson 1983, Wetterer, un- published). In large A. cephalotes colonies, workers primarily attack the leaves and flowers of trees (Cherrett 1968, 1972; Rockwood & Hubbell 1987; Vasconcelos 1990; Wetterer 1994b). These colonies produce an extremely broad continuous size-range of workers (Stradling 1978; Wilson 1983). A wide range of medium-size workers forage, with the larger of these cutting thicker and tougher vegetation (Nichols-Orians & Schultz 1989; Wetterer 1994b). Recent taxonomic research, based on morphological characters and allozyme poly- morphisms, has raised questions as to whether the A. octospinosus on the Pacific and Atlantic slopes of Costa Rica are a single species (J. Longino, T. Schultz, & K. Boomsma, personal communication). Because earlier studies indicated marked inter- specific ecological differences among leaf-cutting ants, we compared the foraging and nesting ecology of Pacific and Atlantic A. octospinosus. We also examined the question as to why A. octospinosus was the only leaf-cutting ant species present at our study site in Palo Verde National Park. METHODS We studied foraging selectivity and nesting habits of Acromyrmex octospinosus col- onies at the start of the wet season (June 1996) in tropical dry forest of Palo Verde Na- tional Park, Guanacaste Province, Costa Rica. We worked in the area around Palo Verde Biological Station (10°19’N, 85°21’W; elevation 10 m), operated by the Organi- zation for Tropical Studies (see Janzen 1983 for a site description). We conducted our study during daylight hours, when most A. octospinosus colonies were actively forag- ing. On an earlier visit to Palo Verde in the dry season (January 1996), there was no diurnal foraging by A. octospinosus (J. Wetterer, personal observation). Through extensive searches along hiking trails and roads around Palo Verde, we located seventeen A. octospinosus colonies. At a nearby open pasture site, Sitio Mojal, we collected data on one additional A. octospinosus colony (colony 12), making a total of eighteen colonies. We found no other leaf-cutting ant colonies at the site, although we did find a single live alate Atta cephalotes queen. We explored the area for several kilometers in different directions, but never located any A. cephalotes colonies. At each A. octospinosus colony, we collected laden foragers from along their forag- ing trails. To help ensure a representative sample, we collected laden foragers at ten second intervals as they passed a designated point on their trail, or if there were few foragers, we collected consecutive foragers. We used data from twelve colonies in our 64 Florida Entomologist 81(1) March, 1998 foraging selectivity analysis. At four large A. octospinosus colonies, we collected 25 for- agers from each of the two main trails. From six smaller colonies, we collected 25 for- agers for each colony. At two very small colonies, we collected only fourteen workers from one colony and eleven from the other, because no more foragers came in a period of several hours. At six other small colonies we collected none or only one forager due to lulls in foraging activity that seemed to be associated with hot sunny weather. We did not include these collections in our foraging selectivity analysis. We classified each load as fresh (soft, pliant, and green) or fallen (dry or yellowed) leaf material (fragment or whole leaf), herb section (stem or stem with leaves, leaf buds, or flowers), flower (part or whole), fruit (part or whole berry), insect frass, or “other.” Whenever possible, we located and identified the original source of each load.
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