Comparison of frugivory by howling monkeys (Alouatta palliata) and bats (Artibeus jamaicensis) in the tropical rain forest of Los Tuxtlas,Mexico

Information on the fruit diets of howling monkeys and fruit-eating bats in the tropical rain forest of Los Tuxtlas, Mexico was collected for a year to compare the plant species used. Howling monkeys used 19 plant species whereas bats used 32 plant species as fruit sources. Eleven species were common in the diet of both mammals. A rank analysis at the plant species level showed that the fruit diets of Alouatta and Artibeus were very different. In contrast to bats, howling monkeys displayed a seasonal pattern in fruit consumption. Diet overlap between the two mammals was highest during the monkeys' fruit-eating season. Measures of fruit production in eight trees (four species) indicated marked variations in fruit biomass produced and in length of fruiting from tree to tree and species to species. Peaks in fruit production were typical both at the species and the individual tree levels, demonstrating the very patchy nature of the fruit available to the monkeys and the bats.     

Leaf-cutting ant preferences for five native tropical plantation tree species growing under different light conditions

To examine the susceptibility of five Costa Rican tree species to leaf-cutter ants (Atta cephalotes L.: Formicidae, Attini), young and mature leaves from trees that were grown in a plantation under full sun and partial shade were offered to six leaf-cutter colonies located in full sun and six in patial shade. In addition to offering leaf disks to the ants, we offered large pieces of leaves to assess the effect of cutting leaves on food choices. Leaf-cutters responded differently to each plant species, preferring Virola koschyni Warburg (Myristicaceae) and Hyeronima alchorneoides Allemao (Euphorbiaceae) over Stryphnodendrum microstachyum Poeppig & Endlicher (Mimosoideae), Pentaclethra macroloba Willdenow (Fabaceae) and Vochysia ferruginea Martius (Vochysiaceae). In agreement with previous studies, interspecific differences among the tree species in water and saponin content appeared to account for the observed ant preferences among the five tree species: leaf-cutter ants preferred leaves and disks with more water and less saponins and the five tree species varied significantly in these traits. An observed positive correlation between preference and phenolic/nitrogen ratio appears to be spurious, and is instead due to a negative correlation between water content and nitrogen content. For the first time, Atta nest location has been shown to affect consumption: nests located in partial shade removed more leaf material than those from the sun plots. However, nest location had no effect on preference ranking of the tree species tested. Pieces of leaves from all the tree species grown in partial shade were significantly more removed than those trees grown in full sun. Leaf age, toughness, leaf specific weights, ant activity, and colony were not correlated with food choices. Because of the observed preferences, the five tree species should not be considered as equal candidates for plantation purposes. However, complete characterization of the candidate status of tree species for plantations in the neotropics must include information on the ability of such species to tolerate pest attacks in addition to their natural defenses to attack.     

Seasonal variation in the positional behavior of red howling monkeys (Alouatta seniculus)

Recent studies on the positional behavior of primates reveal that significant seasonal variation occurs in both locomotion and postures that is related to changes in diet and foraging techniques. Howling monkeys (genusAlouatta), which also have a seasonally varied diet, are predicted to have correspondingly varied positional behaviors. Two groups of red howling monkeys were studied in a primary rain forest in French Guinana during the dry and wet seasons. During the dry season, when howler diet is based mainly on leaves, howlers traveled more frequently by quadrupedal walking on large supports, a mode of progression that is probably inexpensive energetically and relatively stable. During feeding, quadrupedal and tripedal stand contributed considerably, a posture probably associated with the equal distribution of leaves within a tree crown. In contrast, during the wet season, when fruit was abundant, howlers fed very frequently by sitting on large supports, probably because fruit consumption required more time for special manipulation. However, most seasonal changes in feeding postures, and in travel and feeding locomotion, were difficult to associate directly with dietary shifts. These behavioral changes may be more highly correlated with slight modifications in microhabitat use (horizontal and vertical daily ranges, similar and alternative arboreal pathways) that are not considered in this paper.     

Combat between large derived societies: A subterranean army ant established as a predator of mature leaf-cutting ant colonies

Summary. Mature colonies of Atta leaf-cutting ants are dominant herbivores throughout the Neotropics. Although young colonies have natural enemies, mature colonies, which live in extensive nests containing millions of workers, currently have no recognised predators. New World army ants (Ecitoninae) are specialist social predators of other ants, and the army ant Nomamyrmex esenbeckii, a primarily subterranean species, is known to prey upon young Atta colonies. Here we present the results of the first long-term study of the predator-prey interaction between N. esenbeckii and Atta. Our study establishes the army ant N. esenbeckii as the only known predator capable of successfully attacking and killing mature as well as young colonies of Atta leaf-cutting ants. In natural raids, and experimental tests, Atta rapidly recruited their largest workers (majors) as a specific defensive response to N. esenbeckii raiders and both taxa used their largest individuals in the frontline of battles. The deployment and behaviour of these large workers demonstrates a size-related division of labour and agrees with the predictions of Lanchesters Linear Law of Combat. Both taxa also used cooperative combat teams to overwhelm large combatants from the other side. The success of N. esenbeckii raids varied greatly, such that they were prevented from entering Atta nests in the least successful raids, and completely overran Atta colonies in the most successful raids. The speed and magnitude of the defensive response of mature Atta colonies was key in determining the level of success of N. esenbeckii raids.Received 12 December 2003; revised 25 March 2004; accepted 1 April 2004.Work conducted at the Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Ancon, Republic of Panama     

Fruit eating and seed dispersal by howling monkeys (Alouatta palliata) in the tropical rain forest of Los Tuxtlas,Mexico

The frugivory and ranging habits of howling monkeys living in the rain forest of Los Tuxtlas, Veracruz, Mexico, were studied for 5 consecutive years with the aim of investigating the role of seed dispersal agents played by the howling monkeys. The howling monkeys' consumption of fruit occupied half of their feeding time, and 80% of this time was spent feeding on mature fruit. Observations confirmed use of 19 tree species and a preference for a few species of Moraceae and Lauraceae. Fruit exploitation was very seasonal; only two species provided fruit year-round. Home range size was about 60 ha, and animals in the troop (N = 16) showed a day range of 10–893 m. Monthly collection of fecal samples during 2 years indicated that 90% of the seeds (N = 7,928) in the samples belonged to eight tree species and to one liana; the rest belonged to 15 unidentified species of vines. Germination success for the seeds in the feces was about 60% and for control seeds was 35%. Howling monkeys created diverse seed shadows in the vicinity of their leaf and fruit sources, and while they dispersed the seeds of some plant species, they also produced a great deal of fruit and seed waste for others.     

Differences in Forage-Acquisition and Fungal Enzyme Activity Contribute to Niche Segregation in Panamanian Leaf-Cutting Ants

The genera Atta and Acromyrmex are often grouped as leaf-cutting ants for pest management assessments and ecological surveys, although their mature colony sizes and foraging niches may differ substantially. Few studies have addressed such interspecific differences at the same site, which prompted us to conduct a comparative study across six sympatric leaf-cutting ant species in Central Panama. We show that foraging rates during the transition between dry and wet season differ about 60 fold between genera, but are relatively constant across species within genera. These differences appear to match overall differences in colony size, especially when Atta workers that return to their nests without leaves are assumed to carry liquid food. We confirm that Panamanian Atta specialize primarily on tree-leaves whereas Acromyrmex focus on collecting flowers and herbal leaves and that species within genera are similar in these overall foraging strategies. Species within genera tended to be spaced out over the three habitat categories that we distinguished (forest, forest edge, open grassland), but each of these habitats normally had only a single predominant Atta and Acromyrmex species. We measured activities of twelve fungus garden decomposition enzymes, belonging to the amylases, cellulases, hemicellulases, pectinases and proteinases, and show that average enzyme activity per unit of fungal mass in Atta gardens is lower than in Acromyrmex gardens. Expression profiles of fungal enzymes in Atta also appeared to be more specialized than in Acromyrmex, possibly reflecting variation in forage material. Our results suggest that species- and genus-level identities of leaf-cutting ants and habitat-specific foraging profiles may give predictable differences in the expression of fungal genes coding for decomposition enzymes.     

The effect of wilting on the selection of leaves by the leaf-cutting ant Atta laevigata

Using field assays of leaf preference, we tested the hypothesis that wilting affects the selection of leaves by the leaf-cutting ant Atta laevigata (Fr. Smith). Detached leaves were left to air-dry until noticeably wilted. The area removed by the ants from wilted leaves was significantly greater than the area removed from fresh leaves, this effect being observed in several plant species, in leaves of different age, and in assays with different ant colonies. Leaves collected from water-stressed plants were also preferred to leaves from non-stressed plants. A. laevigata was found to employ a two-stage, size-related, strategy when cutting plants. Larger workers climbed the plant stem and dropped whole leaves to the ground by severing their petioles; smaller workers cut the lamina of the dropped leaves. The ants frequently left dropped leaves on the ground, until the next foraging day or even later, when they were harvested in a wilted condition in preference to newly-dropped leaves.It is possible that during wilting some repellent substances evaporate or become less effective, thus enhancing leaf palatability. Alternatively or in addition, changes in nutrient and water content may have rendered wilted leaves more palatable to leaf-cutting ants.     

Host-plant selection,diet diversity,and optimal foraging in a tropical leafcutting ant

Summary A month-long study was conducted on the comparative foraging behavior of 20 colonies of the leafcutting ant, Atta cephalotes L. in Santa Rosa National Park, Guanacaste Province, Costa Rica. The study was conducted during the middle of the wet season, when trees had mature foliage and the ants were maximally selective among species of potential host plants. The colonies always gathered leaves from more than a single tree species but on average one species constituted almost half the diet with the remaining species being of geometrically decreasing importance. Colonies exhibited greater diversity in their choice of leaves and lower constancy of foraging when the average quality of resource trees was lower, as predicted by elementary optimal foraging theory. Furthermore, the ants were more selective of the species they attacked at greater distances from the nest. However, the ants sometimes did not attack apparently palatable species, and often did not attack nearby individuals of species they were exploiting at greater distances.A classical explanation for why leafcutting ants exploit distant host trees when apparently equally good trees are nearer, is that the ants are pursuing a strategy of conserving resources to avoid long-term overgrazing pressure on nearby trees. We prefer a simpler hypothesis: (1) Trees of exploited species exhibit individual variation in the acceptability of their leaves to the ants. (2) The abundance of a species will generally increase with area and radial distance from the nest, so the probability that at least one tree of the species will be acceptable to the ants also increases with distance. (3) The ants forage using a system of trunk-trails cleared of leaf litter, which significantly reduces their travel time to previously discovered, high-quality resource trees (by a factor of 4- to 10-fold). (4) Foragers are unware of the total pool of resources available to the colony. Therefore once scouts have chanced upon a tree which is acceptable, the colony will concentrate on harvesting from that tree rather than searching for additional sources of leaves distant from the established trail.     

Hitchhiking and the removal of microbial contaminants by the leaf‐cutting ant Atta colombica

1. The ecologically dominant leaf‐cutting ants exhibit one of the most complex forms of morphological caste‐based division of labour in order to efficiently conduct tasks, ranging from harvesting fresh leaf material to caring for the vulnerable fungal crop they farm as food. While much of their division of labour is well known, the role of the smallest workers on foraging trails is puzzling. Frequently these minim workers hitchhike on leaf fragments and it has been suggested that they may act to reduce the microbial contamination of leaf material before they enter the nest. Here we investigated this potentially important role of minims with field colonies of Atta colombica. 2. We experimentally increased the microbial load of leaf fragments and found that this resulted in minims hitchhiking on leaf fragments for longer. Furthermore, we show that leaves naturally have a significant microbial load and that the presence of hitchhikers reduces the microbial load of both experimentally manipulated and natural leaf fragments. 3. Intriguingly, the microbial load of leaves high in the canopy where ants were foraging was much lower than closer to the ground where the ants avoided cutting leaves. This suggests that the often perplexing foraging patterns of leaf‐cutting ants may in part be explained by the ants avoiding leaves that are more heavily contaminated with microbes. 4. The removal of microbial contaminants is therefore an important role of hitchhiking minim workers in natural colonies of Atta leaf‐cutting ants, although other tasks such as trail maintenance and defence also explain their occurrence on trails.     

A preliminary study of resource overlap between howling monkeys (Alouatta palliata) and other arboreal mammals in the tropical rain forest of Los Tuxtlas,Mexico

Potential resource overlap between howling monkeys and other arboreal mammals was studied in the rain forest of Los Tuxtlas, Veracruz, Mexico. Eight species of mammals belonging to the orders Primates, Carnivora, Rodentia, and Marsupialia were found to share the canopy and to overlap trophically with howling monkeys. These mammals made up 77% and Alouatta 23% of the arboreal mammalian biomass under consideration. The arboreal porcupine and spider monkey were the only mammals that also fed on leaves. However, in this feeding niche, Alouatta is the only important mammalian folivore in Los Tuxtlas, and resource depression derived from leaf-eating insects is more important. The eight arboreal mammals may exert more pressure upon fruit resources, for they consumed 75% of the estimated total dry weight of fruit/ha/yr consumed by arboreal mammals.     

Howling on the edge: Mantled howler monkey (Alouatta palliata) howling behaviour and anthropogenic edge effects in a fragmented tropical rainforest in Costa Rica

The function of long calling is a subject of interest across animal behaviour study, particularly within primatology. Many primate species have male‐specific long‐distance calls, including platyrrhines like the folivorous howler monkey (Alouatta spp.). Howler monkeys may howl to defend resources such as feeding trees or areas of rich vegetation from other monkey groups. This study tests the ecological resource defence hypothesis for howling behaviour in the mantled howler monkey (Alouatta palliata) and investigates how anthropogenic forest fragmentation may influence howling behaviour. More specifically, this study examines how howling bout rate, duration, precursors and tree species richness, DBH, and canopy cover vary in 100 m anthropogenic edge and interior forest zones at La Suerte Biological Research Station (LSBRS), a fragmented tropical rainforest in Costa Rica. Results show that tree species richness and canopy cover are higher in forest interior at this site, suggesting that monkeys should howl at greater rates in the interior to defend access to these higher‐quality vegetation resources. Overall, our results supported the ecological resource defence hypothesis. The main howl precursor was howling from neighbouring groups. Although howling rate did not differ between forest zones, howling bouts from forest interior were longer, had a greater number of howls per bout and were preceded by different precursors than howls from anthropogenic edge zones, including more howls from neighbouring groups. Our findings provide some of the first evidence for behavioural edge effects in primate vocal communication behaviour.     

Patterns of Foraging of the Forest Ant Formica neogagates Emery (Hymenoptera: Formicidae) on Tree Branches

Numbers of ants moving on trunks of oak trees and successive visits of workers of the forest ant Formica neogagates Emery to leaves of black birch trees were recorded in the forest. Ants were found to search systematically because they tended to move in the same direction on a twig before and after visiting a leaf. However, many leaves were skipped, and the probability of visiting an encountered leaf was about 0.5. Data were used to develop a computer model of ant foraging on leaves and twigs of trees. It was found that reasonable model outputs of numbers of leaves skipped between visits and the number of new leaves visited, as the total number of visits increased, could be obtained only if ants were assumed to partially avoid leaves that they had previously visited. Model results implied that 100 ants in a tree foraging for about 3 h should be able to visit about one quarter of the leaves in a tree.     

Look before leaping: foraging selectivity of capuchin monkeys on acacia trees in Costa Rica

Acacia trees in Costa Rica have an obligate mutualism with three species of Pseudomyrmex ants, which vigorously defend their host tree from insect and mammalian herbivores. Depending on the size and species of ant colony, individual acacia trees may be differentially protected. For animals able to discern between weakly and highly aggressive ant colonies, costs of ant stings from less active colonies might be offset by nutritional value acquired from feeding on acacia fruit or ant larvae in swollen thorns. We examined foraging selectivity of capuchin monkeys on acacia trees in Santa Rosa National Park, Costa Rica. We measured four characteristics of the acacia trees from which capuchins fed and of acacias immediately adjacent to those in which the monkeys fed: diameter at breast height (DBH), accessibility, species of closest tree and ant species present. We found that capuchins prefer to forage in acacias that are large and accessible. We also made two measurements of ant colony activity on each tree, one before and one after disturbing the ant colony. We found that the three species of mutualistic ants differ in baseline activity levels and that mutualistic ants are more active than non-mutualistic ant species found in acacia trees. We also found that capuchins foraged more frequently in trees colonized by non-mutualistic ants, but the explanatory value (r ²) of this model was low. Furthermore, monkeys did not discriminate between acacias on the basis of baseline ant activity or the ant colony’s response to disturbance. We conclude that these monkeys select acacia trees in which to forage based on characteristics of the trees rather than the ants. In addition, our study suggests that white-faced capuchins act as predators on the acacia ants but they probably benefit the dispersal and reproductive success of acacia trees. Capuchins may in fact function as an additional mutualistic partner for acacia trees via seed dispersal, but they must overcome the ants’ defense of the trees to do so.     

Foraging Behavior and Activity Patterns of Alouatta caraya in the Northeastern Argentinean Flooded Forest

Howling monkeys (Alouatta spp.) are colonizer species, showing a plasticity in behavior that allows them to inhabit different sorts of forests. There is a series of hypotheses relating demography to behavioral and ecological characteristics of howlers: (1) as howler density increases, home range size will decrease; (2) howler groups occupying small home ranges will have a high proportion of leaves in their diet; and, (3) the proportion of mature leaves in the diet is negatively correlated with group daily travel distance. To test hypotheses about howler diet and ranging pattern in relation to food resources, we studied the foraging ecology and general activity patterns of 2 groups of black howlers (Alouatta caraya) inhabiting in flooded forest of Paraná River (Argentina), the habitat with the highest density recorded for Alouatta. We found, in addition to the highest densities, also the smallest home ranges for Alouatta (Group I: 1.7 ha and Group II: 2.2 ha). However, diet was not dominated by leaves, and high quality items—flowers, fruits, new leaves—were a high proportion of the diet all year. Also, daily travel distance is not correlated with the proportion of mature leaves in the diet, but is positively correlated with the number of group confrontations. Finally, we propose a model integrating and interpreting our results as a function of the spatial and temporal distribution of high quality food resources in the Argentinean flooded forest.     

Foraging and leaf-cutting of the desert gardening ant Acromyrmex versicolor versicolor (Pergande) (Hymenoptera: Formicidae)

Summary Four colonies of the desert leaf-cutter ant Acromyrmex versicolor versicolor (Pergande) located 30 miles N.E. of Tempe, Arizona were observed over a 7 month period. The ants utilized trails in foraging, a characteristic of higher attines, as well as foraging singly, a typical pattern among the more primitive gardeners. The ants cut and collected both dry and green vegetation with dry grasses comprising the bulk of the forage. The ants increased their cutting of green vegetation after significant rainfall but collected dry grasses almost exclusively during dry periods. Detailed macro-motion picture analysis of leaf-cutting revealed that the desert gardener utilized a unique technique for cutting compound desert leaves.     

Ontogenetic changes in forager polymorphism and foraging ecology in the leaf-cutting ant Atta cephalotes

In the leaf-cutting ant Atta cephalotes (L.) small colonies produce a relatively narrow size-range of small workers, whereas large colonies produce a much wider size-range of workers. In this study, I compared the foraging of four small A. cephalotes colonies (fewer than 5000 workers) with published data on foraging of large colonies to examine how colony size and worker size-range may be related to foraging ecology in leaf-cutting ants. I found that the foraging ecology of small A. cephalotes colonies is very different from that of large colonies. In small colonies, a relatively narrow size-range of foragers (1.4–6.7 mg, mean 3.3 mg) cut primarily herbs (ferns, grasses, and other small herbaceous plants) located within 7 m of the nest. In contrast, in large colonies, a broader size-range of workers (1.4–30 mg, mean 7.3 mg) participate in foraging, generally harvesting from trees 20–80 m from the nest, with larger workers cutting on trees with thicker and tougher leaves. Small colonies' dependence on small herbaceous plants near the nest may have a profound impact on distribution of A. cephalotes. A. cephalotes colonies are rarely found in primary forest, where the low occurrence of small herbaceous plants in the understory may preclude the establishment of young colonies.     

Habitat selection and seasonal patterns of activity and foraging of mantled howling monkeys(Alouatta Palliata) in Northeastern Costa Rica

I conducted a 15- month ecological study of habitat preferences and activity and foraging patterns of two troops of mantled howling monkeys, Alouatta palliata,in a lowland rain forest at La Selva Biological Reserve in northeastern Costa Rica. The two troops specialized on different habitats in spite of the fact that both of them had all habitats available and were not constrained by neighboring troops since the population density of howlers is low (7- 15 howlers/km ² ).Troop 1 spent the majority of time in primary forest (80%) followed by secondary forest (10%), while troop 2 spent the majority of time in undisturbed riparian habitat (60%) followed by primary forest (30%). Habitat sampling indicates that neither the total number of stems, species, or families nor the diversity (Shannon index) or evenness is a good indicator for howler habitat selection. Instead the density of trees from the 12 species most commonly consumed by each troop is the most important factor. Activity and foraging patterns were not dependent upon the season as has been described for howling monkeys in forests with a more pronounced dry season at Barro Colorado Island, Panama, and La Pacifica in northwestern Costa Rica. This is likely a result of the more constant food supply at La Selva, combined with less intraspecific competition due to the low howler density. The intraspecific variability of foraging patterns and troop- specific habitat specialization observed in Alouatta palliatashould be considered in the conservation biology of primates. Primate relocation programs should include not only an ecological assessment of the release site but also a comparison of the release site with the habitat that the groups currently occupy.     

Leaf nutrient content and host plant selection by leaf-cutter ants, Atta laevigata, in a Neotropical savanna

The selection of nutrient-rich leaves by leaf-cutter ants ( Atta spp.) is thought to be of indirect benefit to these ants by promoting the growth of their symbiotic fungus. However, relatively few studies have analyzed the influence of leaf nutrient content on host plant selection by leaf-cutter ants, and conflicting results have been found. We compared the content of eight nutritional elements plus the non-nutrient aluminum between leaves harvested by colonies of Atta laevigata (Smith) (Hymenoptera: Formicidae: Attini) and leaves collected randomly within their foraging areas. In addition, we evaluated whether leaf nutrient content explained the frequency with which these ants attacked and defoliated some of the tree species found in the study area. For 2 years, we monitored 17–26 trees from 15 species and determined the number of times each plant was attacked and the amount and type of foliage removed. Leaves harvested by A. laevigata presented significantly higher concentrations of N, P, K, Zn, and Cu than those collected randomly. This result is likely to reflect the foraging pattern presented by these ants, which were selective both in terms of the plant species and age of leaves most commonly attacked. Young leaves were the only or the main leaf type exploited in many species, and in comparison to mature leaves these presented significantly higher concentrations of P and K. Large differences in the mean number of ant attacks on the tree species studied were also observed, and those presenting more leaf N tended to be the most frequently attacked.     

Performance and fate of tree seedlings on and around nests of the leaf‐cutting ant Atta cephalotes: Ecological filters in a fragmented forest

Habitat fragmentation is currently the most pervasive anthropogenic disturbance in tropical forests and some species of leaf‐cutting ants of the genus Atta (dominant herbivores in the neotropics) have become hyper‐abundant in forest edges where their nests directly impact up to 6% of the forest area. Yet, their impacts on the regeneration dynamics of fragmented forests remain poorly investigated. Here we examine the potential of Atta cephalotes nests to function as ecological filters impacting tree recruitment. Growth, survival and biomass partitioning of experimentally planted seedlings (six tree species) were examined at eight spatially independent A. cephalotes colonies in a large Atlantic Forest fragment. Seedling performance and fate (leaf numbers and damage) were monitored up to 27 months across three habitats (nest centre, nest edge and forest understorey). Plants at illuminated nest centres showed twice the gross leaf gain as understorey individuals. Simultaneously, seedlings of all species lost many more leaves at nests than in the forest understorey, causing a negative net leaf gain. Net leaf gain in the shaded understorey ranged from zero (Licania and Thyrsodium species) to substantial growth for Copaifera and Virola, and intermediate levels little above zero for Protium and Pouteria. Also seedling survival differed across habitats and species, being typically low in the centre and at the edge of nests where seedlings were often completely defoliated by the ants. Lastly, seedling survival increased strongly with seed size at nest edges while there was no such correlation in the forest. Our results suggest that Atta nests operate as ecological filters by creating a specific disturbance regime that differs from other disturbances in tropical forests. Apparently, Atta nests favour large‐seeded tree species with resprouting abilities and the potential to profit from a moderate, nest‐mediated increase in light availability.     

Patterns of colonization and incipient nest survival inAcromyrmex niger andAcromyrmex balzani (Hymenoptera: Formicidae)

Summary Patterns of colonization by queens and incipient nest survival of the leaf-cutting antsAcromyrmex niger andAcromyrmex balzani were studied by means of belt transects and individually marked incipient nests. No relation was found between colony density and the number of colonization attempts. Both species are not claustral, and high rates of queen mortality were attributed to conspecific executions and predation. Of other discernable mortality factors, failure of fungal garden establishment was the most important. Only 34 of 296A. balzani and 13 of 154A. niger marked colonies were alive at the end of one year. These figures are higher than those reported for species ofAtta. These results are contrasted with those of claustral-foundingAtta species. Small colonies are occasionally raided by larger colonies which robbed brood.