Pogonomyrmex Barbatus
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Anita. Behav., 1986, 34, 1402-1419 The dynamics of the daily round of the harvester ant colony (Pogonomyrmex barbatus) DEBORAH M. GORDON Museum of Comparative Zoology and Society of Fellows, Harvard University, Cambridge, Massachusetts 02138, U.S.A. Abstract. Colonies of the red harvester ant, Pogonomyrmex barbatus, do various tasks outside the nest. There is a daily temporal pattern in the numbers of ants engaged in each of five activities: foraging, nest maintenance, patrolling, midden work and convening. Perturbations were carried out in the field to investigate how the daily round changes in response to environmental events and colony needs. Interfering with nest maintenance, foraging or both caused changes in the temporal patterns in all five of the observed activities. Removing nest maintenance workers, foragers or both caused the numbers involved in all five activities to decrease, and there were temporal patterns in the effects of removals. The results of both interference and removal experiments show that the extent to which a worker group does one activity affects the behaviour of other groups. When nest maintenance or foraging is impeded experimentally, these two activities are of reciprocal priority. When both are impeded, foraging is of higher priority than nest maintenance. Harvester ants forage, patrol, maintain the nest speaking, that of an equilibrium state, but the area and foraging trails, collect and arrange dynamics of colony organization are not yet under- pebbles on the nest, and gather in small groups, stood. What events will alter the daily round, by inspecting and grooming each other. The beha- how much, and for how long? Is there more than viour of the colony outside the nest at any moment one equilibrium state? can be described by citing the number of ants This study begins an exploration of the dynamics engaged in each of the five activities. The numbers relating colony priorities and external events. I change throughout the day in a predictable man- perturbed the daily rounds of harvester ant col- ner. There is a daily temporal pattern in the onies and examined the resulting changes in daily numbers of ants engaged in each activity. Describ- rounds. I did two kinds of perturbation experi- ing the pattern in all activities is describing the daily ments: first, interfering with nest maintenance, round of the colony. foraging, or both; and second, removing nest Daily rounds are colony-specific (Gordon maintenance workers, foragers or both. In the 1983a) and species-specific (Gordon 1984a), and interference experiments, nest maintenance response to experimental manipulations may workers were presented with a pile of toothpicks depend on the stage in the daily round when that had to be cleared away before other activities experiments are done (e.g. Gordon 1983b). Thus, could be done and barriers were placed across the the daily round of a harvester ant colony provides a foraging trails to prevent foragers from going baseline from which to investigate colony beha- directly to their destination. In the removal experi- viour. Certain aspects of chemical communication, ments, workers engaged in each task were removed response to new food sources and territorial beha- from the colony. viour can be explained in the context of the daily The following questions were considered. round (Gordon 1983b,c, 1984b). (1) How independent are different worker It would be a mistake, however, to think of the groups? First, do changes in the temporal pattern daily round as a rigid programme. Temporal of one activity affect temporal patterns in other patterns in the numbers of ants engaged in each activities? Little is knownabout the dynamics of activity can change. On some days, for example, the temporal relationships between the different foraging predominates; on others, colonies do tasks of an ant colony. By interfering with nest mainly nest maintenance work (personal observa- maintenance and foraging, I changed the temporal tion). The concept of the daily round is, loosely pattern in each of these two activities. Subsequent 1402 Gordon: Dynamics of daily round of harvester ants 1403 temporal patterns in other activities were then INTERFERENCE EXPERIMENTS examined. Second, does the removal of ants doing nest maintenance and/or foraging affect the Methods numbers of workers, and temporal patterns, in The study was conducted in a chapparal-mes- other activities? Individual ants are known to quite habitat in the Lower Sonoran desert near switch tasks in response to environmental distur- Rodeo, New Mexico, during a stay at the South- bances (Meudec & Lenoir 1982) and the removal of western Research Station. Twelve colonies of the workers (Wilson 1983, 1984). Here I consider how, harvester ant Pogonomyrmex barbatus were at the group level, a decrease in the numbers observed during six observation periods from 19 available to do one task affects the numbers and July to 30 August 1984. All observation periods temporal patterns in other tasks. (OP) were 6 days long, except for OP5, which was (2) Which aspects of the daily round are flexible? only 3 days long. Each colony was observed once First, are nest maintenance and foraging of equal each hour from 0600 until 1300 hours. All activities priority? Common sense suggests that foraging is seen outside the nest were classified into the five of a high priority to an ant colony. However, activity types listed above and in Table I. An harvester ant colonies store seeds inside the nest observation consisted of recording the numbers of (MacKay 1981) and are able to survive periods of ants engaged in each activity within 1.3 m of the inactivity (Whitford & Ettershank 1975). Nest nest entrance. This area contains workers in all five maintenance workers clear and maintain the forag- activities, including some of the foragers, but only ing trails and nest mounds, which are thought to be about the first 0.3 m of the foraging trail. The sum the site of colony-specific pheromones used in of the workers engaged in the five activities repre- territorial behaviour (Holldobler 1976). Thus nest sents the total number of ants outside the nest. maintenance may be crucial in interspecific and Experiments were conducted during OP2, OP4, inter-colony relationships, as well as in keeping the and OP6. Six of the 12 colonies were left undis- nest mound clear for other activities, and is con- turbed as controls. The other six were divided into ceivably as important to the colony, in the short three groups of two colonies each. In one group, a term, as foraging. pile of toothpicks was placed near the nest entrance If foraging and nest maintenance are of equal after the 0700 hours and before the 0900 hours priority, then perturbing each activity singly would observations, when at least nine maintenance have similar effects, and perturbing both would workers had been seen at work on the nest mound. have the same effect, but of larger magnitude. The number of toothpicks was increased as follows: Second, which aspects of the daily round are OP2, 100 toothpicks; OP4, 200 toothpicks; OP6, preserved despite perturbations? Patrolling pre- 300 toothpicks. The ants usually moved all the cedes foraging in the daily round of undisturbed toothpicks to the perimeter of the nest mound colonies, and it appears that patrollers recruit within 1 h. Once there, the toothpicks were no foragers to new food sources (Gordon 1983b). If longer manipulated by the ants. I removed all the the temporal pattern in foraging is changed, will toothpicks at the end of each day. patrolling still precede it? I examined which tem- In the second group of two colonies, plastic poral relationships between activities persist after rectangular barriers (15x 10 cm) with two V- perturbations. shaped indentations (4 cm long) cut, wide end The methods used in this study reflect a concern down, into the bottom of each rectangle, were with patterns in group, rather than individual, placed across foraging trails. Each rectangle was behaviour. The daily round is an example of such a bent to a 90 ~ angle lengthwise, across the bases of pattern. In exploring whether and how temporal the Vs, and held on the ground with rocks. Ants patterns in different activities are related to each approaching from one side encountered the wide other, this study has two goals. The first is to end of the V and passed through the barrier easily. examine the relationship between colony priorities From the other side, they encountered only a small and external events in a harvester ant colony. The opening, formed by the base of the V. The presence second is to offer some tractable questions about of the barrier caused interference on the foraging group behaviour that may be useful in investigating trail, as the ants went either back and forth along it the organization of other animal societies. until getting through an opening, or around the 1404 Animal Behaviour, 34, 5 .~ ~ ~ o .~ ~ ~ ~ ~ ~ ~ ~ ~o~ o o o 0 Gordon: Dynamics of daily round of harvester ants 1405 barrier. The number of barriers was increased as ment with the undisturbed colonies. Thus the follows: OP2, two barriers on one trail; OP4 and undisturbed colonies in the same observation OP6, four barriers on two trails (the number of period were used as controls. barriers was not increased in OP6 since most To test whether experimental and undisturbed colonies had only two major trunk trails). Barriers colonies were alike before the experiments, a were placed 1-5 m and 1.8 m from the nest entrance, similar ANOVA was done on the data from OP1 (a outside the 1-3 m radius of observation, and three-way ANOVA since there was no OP effect). oriented so that ants going in either direction along To test for residual effects of the experiments, the trail would encounter the wide and narrow similar ANOVAs were done using the data from openings once each.