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Seasonal Patterns in Activity Rhythm and Foraging Ecology in the Neotropical Forest-Dwelling Ant, Odontomachus Chelifer (Formicidae: Ponerinae)

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Seasonal Patterns in Activity Rhythm and Foraging Ecology in the Neotropical Forest-Dwelling Ant, Odontomachus Chelifer (Formicidae: Ponerinae) BEHAVIOR Seasonal Patterns in Activity Rhythm and Foraging Ecology in the Neotropical Forest-Dwelling Ant, Odontomachus chelifer (Formicidae: Ponerinae) 1 RAFAEL L. G. RAIMUNDO, ANDRE´ V. L. FREITAS, AND PAULO S. OLIVEIRA Departamento de Biologia Animal, Universidade Estadual de Campinas, C.P. 6109,13083-862 Campinas SP, Brazil Ann. Entomol. Soc. Am. 102(6): 1151Ð1157 (2009) ABSTRACT We provide qualitative and quantitative data on the natural history and foraging behavior of the ground-dwelling ant Odontomachus chelifer (Latreille) (Formicidae: Ponerinae) in a forest reserve in southeastern Brazil, with emphasis on colony activity rhythms and diet preferences in relation to seasonal availability of potential food items in the leaf litter. Ant colonies exhibited nocturnal activity throughout the year, and they foraged signiÞcantly more intensively in the wet/ warm (NovemberÐMarch) than in the cold/dry season (AprilÐOctober). As the night begins, small groups of workers disperse and hunt individually on a wide diversity of litter arthropods of variable sizes. At dusk, encounters with foragers of the diurnal ponerine Pachycondyla striata Fr. Smith were conspicuously avoided by O. chelifer, which occasionally had their prey robbed by the former or were even taken as prey themselves. Termites were the preferred prey of O. chelifer, making up 40% of the food items captured in each season. Seasonal comparisons of prey organisms captured by O. chelifer, and of litter-dwelling arthropods sampled in pitfall traps, revealed that the frequency distribution of retrieved prey in each taxonomic group did not differ seasonally, despite the 2.7-fold increase in the overall availability of litter arthropods in the warm/wet period. This result supports foraging theory by showing that preference for certain animal prey types (i.e., taxonomic groups) persists through time despite seasonal ßuctuations in the overall availability of potential prey on the forest ßoor. This study points out to the importance of studying ant foraging ecology and diet preferences in a natural context. KEY WORDS activity schedule, diet, ground-dwelling ants, ant behavior The daily activity schedule is one of the most distinc- Nonetheless, a diversity of feeding habits and foraging tive characteristics among species of ants (Ho¨lldobler modes have been reported for members of this sub- and Wilson 1990). InterspeciÞc divergence in activity family, which include hunting for food on ground and patterns result from particular morphological, physi- foliage substrates, scavenging for dead arthropods, ological or behavioral traits that deÞne the ecological gathering plant and insect exudates, and collecting tolerance of a species and thus determine its speciÞc ßeshy fruit (Duncan and Crewe 1994, De´jean and foraging period (Bernstein 1979). The temporal di- Suzzoni 1997, Blu¨ thgen et al. 2003, Oliveira and Freitas mension of ant foraging behavior is constrained by 2004, Dutra et al. 2006). In addition, although many several abiotic and biotic factors that might affect the ponerines feed opportunistically on a diversity of food costs of foraging and the use of time (Traniello 1989). types, some specialized species can be extremely spe- Common abiotic factors affecting ant activity sched- ciÞc in the kind of prey they consume (Freitas 1995, ules include temperature (Cerda´ et al. 1998) and mois- Leal and Oliveira 1995). Foraging strategies may range ture (Levings and Windsor 1984). Biotic factors such from solitary to cooperative hunting, with varying as interspeciÞc competition (Carroll and Janzen degrees of recruitment behavior among nestmates 1973), natural enemies (Orr and Seike 1998), and (Peeters and Crewe 1987). variation in resource availability (Briese and Macau- Ant species in the genus Odontomachus are widely ley 1980) also may inßuence activity patterns both on known by their trap-jaws used during hunting, which a daily and seasonal basis. can instantaneously stun or kill prey, or during defense Ants in the subfamily Ponerinae often have small against natural enemies (Carlin and Gladstein 1989). colonies with workers that exhibit solitary foraging When a potential prey contacts the trigger hairs of (Peeters and Crewe 1987). Because ponerine ants are Odontomachus, the trap-jaws may close in Ͻ0.1 ms, armed with a sting and many species have powerful resulting in a powerful mandible strike that has been mandibles, they are usually considered as predators. considered an adaptation to feed on fast and soft- bodied arthropods (Spagna et al. 2008). Solitary for- 1 Corresponding author, e-mail: [email protected]. agers of Odontomachus are facultative termite-hunters 0013-8746/09/1151Ð1157$04.00/0 ᭧ 2009 Entomological Society of America 1152 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 6 (Fowler 1980, De´jean and Bashingwa 1985) that also riod. To avoid disturbance of ant foragers, no food item may feed on a wide range of invertebrate taxa (Ehmer was collected during sessions estimating the daily ac- and Ho¨lldobler 1995) and also explore alternative re- tivity schedule of ant colonies. If possible, in such cases sources such as insect honeydew (Carroll and Janzen the food item was identiÞed visually only. 1973), extraßoral nectar (Blu¨ thgen et al. 2003), and Seasonal differences in the availability of potential lipid- and protein-rich ßeshy seeds and fruit (Pizo and prey consumed by O. chelifer was evaluated with 10 Oliveira 2001, Passos and Oliveira 2003). Because pitfall traps distributed on the ground in areas of thick availability of insect prey (Janzen and Schoener 1968) leaf litter, along a 100-m transect in the forest. Traps and plant-derived resources (Rico-Gray and Oliveira were 10 m apart from one another, and were left open 2007) exploited by ants have a strong seasonal com- from 2000 to 2400 hours during three nonconsecutive ponent, some ant species may change their foraging nights in each season. Traps consisted of plastic bottles patterns and/or dietary preferences throughout the (10 cm in diameter) containing 200 ml of 70% ethanol year to optimize food intake in accordance with col- solution. ony phenology (e.g., Judd 2005). The genus Odontomachus occurs widely in tropical Results and warm temperate regions and is especially abun- dant in the neotropics, ranging from semiarid envi- Nests of O. chelifer are located on the ground, with ronments to rain forests (Brown 2000). This article chambers and galleries extending among roots of ad- presents a detailed Þeld account of the foraging ecol- jacent trees. The single nest entrance measures Ϸ5cm ogy of the ant Odontomachus chelifer (Latreille) (For- in diameter and is usually surrounded by a layer of micidae: Ponerinae), a widely distributed species in humid leaf litter. Worker ants, winged forms, dealated the Neotropical region that occurs in forest habitats queens, and immature stages can be found all the way from Mexico to Argentina (Kempf 1972). We provide down to Ϸ1.0 m below the soil surface. qualitative and quantitative data on the natural history O. chelifer exhibited a distinct nocturnal foraging of O. chelifer, with emphasis on colony activity activity throughout the year. The activity schedule is rhythms and dietary requirements across seasons in a clearly inßuenced by the photoperiod (Fig. 1). Over- forest reserve in southeastern Brazil. More speciÞ- all daily activity was negatively affected by ground ϭϪ ϭ cally, we evaluate diet preferences in relation to the temperature in the wet/warm season (rs 0.65, P ϭϪ ϭ seasonal availability of potential food items in the 0.022), but not in the dry/cold season (rs 0.47, P environment. 0.12). Seasonal variation in the activity of ant colonies Study Site and Methods. Fieldwork was carried out was evident. During the warm/wet season the ants from August 1999 to July 2000 at the Santa Genebra foraged intensively throughout the night, with few forest reserve in Campinas, southeastern Brazil (22Њ workers being observed outside their nests at daytime 49Ј S, 47Њ 06Ј W; 670-m altitude). The vegetation con- (Fig. 1A). However, in the cold/dry season colony sists of a semideciduous Atlantic forest fragment. The foraging activity decreased markedly and very few climate of the region exhibits considerable seasonal ants were seen at night only (Fig. 1B). variation, with a warm/wet season lasting from No- Foragers of O. chelifer searched for food chießy vember to March (highest mean monthly records: above and beneath the leaf litter, and only occasion- rainfall, 240.2 mm; temperature, 24.4ЊC) and a cold/ ally they were seen hunting on herbaceous vegetation. dry season lasting from April to October (lowest mean At sunset, small groups of workers were usually ob- monthly records: rainfall, 36.8 mm; temperature, served motionless at the nest entrance. As the night 18.2ЊC) (Toniato et al. 1998). begins the ants disperse from these groups and forage Ten nests of O. chelifer were tagged in the forest for individually in the midst of fallen leaves. As returning behavioral observations. Focal nests were 10 m off the workers start bringing prey into the nest, ant depar- main trail of the reserve and were at least 10 m away ture from stationary groups at nest entrances tends to from one another. Food items were collected directly increase. As they search for food, foragers keep their from the mandibles of returning foragers from any of trap-jaws open and frequently touch the substrate the colonies. Prey items were preserved in 70% alcohol with the antennae. When a potential prey is found, the and brought to the laboratory for more detailed iden- ant forager approaches cautiously and strikes with the tiÞcation, and measurement of length. Total time mandibles. As a result the prey can be injured and spent sampling food items during cold/dry and warm/ the ant is usually thrown 10Ð15 cm backward. This wet season was 21 and 18 h in each period, respec- behavioral sequence can be repeated many times be- tively.
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