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Territory Defense by the Ant Azteca Trigona: Maintenance of an Arboreal Ant Mosaic Author(S): Eldridge S

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Territory Defense by the Ant Azteca Trigona: Maintenance of an Arboreal Ant Mosaic Author(S): Eldridge S International Association for Ecology Territory Defense by the Ant Azteca trigona: Maintenance of an Arboreal Ant Mosaic Author(s): Eldridge S. Adams Reviewed work(s): Source: Oecologia, Vol. 97, No. 2 (1994), pp. 202-208 Published by: Springer in cooperation with International Association for Ecology Stable URL: http://www.jstor.org/stable/4220605 . Accessed: 21/09/2012 17:16 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Springer and International Association for Ecology are collaborating with JSTOR to digitize, preserve and extend access to Oecologia. http://www.jstor.org Oecologia (1994) 97:202-208 Springer-Verlag 1994 ORIGINAL PAPER Eldridge S. Adams Territory defense by the ant Azteca trigona: maintenance of an arboreal ant mosaic Received: 16 November 1992 / Accepted: 19 November 1993 Abstract Mosaics of exclusive foraging territories, pro- Introduction duced by intra- and interspecific competition, are com- monly reported from arboreal ant communities Ant communities are often highly structured due to in- throughout the tropics and appear to represent a recur- tra- and interspecific competition (H?lldobler and ring feature of community organization. This paper Wilson 1990). The "ant mosaic hypothesis" suggests documents an ant mosaic within mangrove forests of that interspecific competition produces a recurring pat- Panama and examines the behavioral mechanisms by tern of community organization among arboreal ants which one of the common species, Azteca trigona, main- throughout the world's tropics (Leston 1973a, b; Majer tains its territories. Most of the mangrove canopy is 1972,1976a, b; Room 1971, 1975a, b). A central claim of occupied by mutually exclusive territories of the ants A. this hypothesis is that the numerically dominant ant trigona, A. velox, A. instabilis, and Crematogaster bre- species within the forest canopy defend mutually exclu- vispinosa. When foraging workers of A. trigona detect sive foraging territories, which nearly fill available workers of these territorial species, they organize an space. Such territorial ant mosaics have been document- alarm recruitment response using pheromonal and tac- ed among canopy ants in lowland tropical forests in tile displays. Nestmates are attracted over short dis- various regions of the world, especially in agricultural tances by an alarm pheromone originating in the py- settings (Way 1953; Brown 1959; Room 1971, 1975a, b; gidial gland and over longer distances by a trail Majer 1972, 1976a, b; Leston 1973a, 1978; Jackson pheromone produced by the Pavan's gland. Recruits are 1984). simultaneously alerted by a tactile display. No evidence The behavioral mechanisms underlying the forma- was found for chemical marking of the territory. Major tion of territory mosaics are in most cases poorly under- workers are proportionally more abundant at territory stood. Although intra- and interspecific aggression have borders than on foraging trails in the interior of the been observed frequently among tropical arboreal ants colony. The mechanisms of territory defense in A. trigo- (e.g., Way 1953; Brown 1960; Vanderplank 1960; Majer na are remarkably similar to those of ecologically 1976b), there have been few investigations of the mecha- analogous ants in the Old World tropics. nisms by which territory mosaics are produced (but see H?lldobler and Wilson 1978; H?lldobler 1979, 1983; ? ? ? Key words Ants Territoriality Azteca Ph?romones Salzemann and Jaffe 1990). Studies from the New World tropics are particularly uncommon. Colony defense in- volves the coordinated actions of hundreds or thou- sands of workers, linked together by diverse forms of communication (H?lldobler and Lumsden 1980). This paper reports that the mechanisms of communication which allow the maintenance of mosaics are remarkably E. S. Adams1 similar for two distantly related ant genera for which Department of Zoology, University of California, detailed information is available. California USA Berkeley, 94720, This study examined territory defense by the abun- Smithsonian Research Institute, Tropical dant ant Azteca and its APO Miami 34002, USA Neotropical trigona (Emery) congener A. velox (Forel). Field experiments and obser- Present address: vations were conducted within a naturally occurring ant 1 Department of Biology, mosaic to document the nature of the mosaic, to deter- University of Rochester, Rochester, NY 14627, USA mine how these ants organized territory defense, to de- 203 termine the role of caste in defense, and to test for the had been collected was used to assay responses to the gland ex- tracts. A of the donor of A. was collected presence of a territorial pheromone (H?lldobler and fragment colony trigona and transferred to a container, Wilson laboratory measuring approxi- 1978). mately 30 ? 50 ? 20 cm. The colony was provided with a paper- covered foraging arena marked with a grid of lines spaced 3 cm apart. For each test, 10 ?? of fluid containing the contents of one worker's gland were used to mark a randomly selected square. Methods After allowing 1 min for the ethanol solvent to evaporate, the ants within the marked were counted. As a control, Study site and species standing square some squares were marked with ethanol alone. For A. a similar was conducted in the field. In each Studies were conducted in forests near Coco Solo, velox, assay mangrove 10 of the contents of two workers' Colon Province, on the Atlantic coast of the of Panama trial, ?? extract, containing Republic were to a 1 cm2 area of bark near a occu- from 1983 to 1989. characteristics are described by glands, applied densely Vegetation trail. After 1 min for the ethanol to Adams and Levings (1987). Ants were identified by R. Snelling pied foraging waiting evapo- the number of ants near the treated area and main- and vouchers have been deposited at the Los Angeles County rate, standing was counted. Ethanol was used as a Museum of Natural History. Because the taxonomy of Azteca is in taining aggressive postures control. need of revision and Vinson 1989), these identifications (MacKay For statistical data for A. velox were may be subject to change. analysis, log-transformed in order to equalize variances among responses to the four cate- gories of extracts. This was not necessary for data from A. trigona. The T-method was used to evaluate Mapping unplanned comparisons among the means (Sokal and Rohlf 1981). Maps of foraging territories were made near ground level ( < 2m In mangrove forests, ants are usually from height). prevented Glandular source of the trail walking on the ground by tidal flooding. Nonetheless, colonies of pheromone Azteca spp. and other arboreal ants often forage near the water Extracts were as described above. A 25 ? 40 cm surface and rely upon connections near this level to travel from prepared platform was covered with and in the field near a test tree to tree. The lateral extent of each colony's foraging area was paper placed colony of A. or A. velox. A 1-2 m connected the determined by following foraging workers and by searching the trigona bridge platform to a route of the test Ants were baited to base of each tree for activity. When foraging workers were not major foraging colony. one end of the with beef until a of at least 100 discovered during visual scans, baits of tuna or beef were placed platform group workers had collected around the bait. Artificial trails 25 cm in and monitored for 2 hr. Intraspecific boundaries were located for were then drawn across the with a Azteca spp. (but not for a co-occurring territorial ant, Cremato- length paper syringe, leading outward from the bait. of extract the gaster brevispinosa [Mayr]) by transferring workers among baits Twenty ?? containing glands from four workers were used to each trail. Control trails and foraging trails to determine if fighting occurred. The edges of prepare were drawn with 20 of ethanol. each territory were marked with colored flagging and mapped ?? The ants each trail were counted a 3 min from central trees using a compass and tape measure. following during period. For statistical analysis, the reciprocals of counts of A. velox were used in order to equalize variances. Due to the lack of variation in A. to two of the Caste participation response by trigona categories trails, Kruskal-Wallis test was used instead of ANOVA. To compare the caste distribution of ants defending territory bor- ders and ants in other parts of the territory, workers were collect- Territorial ed at different sites from 10 territories of A. trigona. The borders pheromone of neighboring territories are defended by dense groups of work- colonies of A. were chosen. Within the ers, whose behavior differs from that of foraging workers in the Twenty trigona foraging area of each a circle of filter 15 cm in diameter was interior of the territory (Adams 1990). For each colony, workers colony, paper attached to a tree trunk with dense trails of A. were collected by rapidly pressing a strip of tape across a border foraging trigona. The resident ants soon established trails across the disks. defense group so that ants were removed before alarm responses paper The disks were retrieved after ten and the of ants could change the group composition. A group of workers was days responses to three of were measured in the field: collected in the same manner from a major foraging trail in the types paper (1) paper previ- their own interior of the colony. The head width of each ant was measured ously occupied by colony, (2) paper previously occupied a A.
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