Seasonal polydomy in a polygynous supercolony of the odorous house ant,Tapinoma sessile

Abstract 1. The odorous house ant, Tapinoma sessile, is a native ant species common throughout North America. In its natural habitat, T. sessile is a low‐key species that consists of small colonies. In invaded urban areas, T. sessile exhibits extreme polygyny and polydomy and becomes a dominant invasive pest. 2. The current study examined: (i) the density, persistence, and the spatial distribution of nests in a large supercolony of T. sessile, (ii) trail abundance and overall colony connectivity as facilitated by the network of trails, (iii) the abundance and the spatial distribution of competing ant species, and (iv) the effect of environmental factors on the number and distribution of T. sessile nests. 3. A distinct pattern of seasonal polydomy was observed, whereby the colony undergoes an annual fission‐fusion cycle. The colony occupies one or a few nests during the winter, experiences rapid exponential growth in the spring to colonize available nesting sites, reaches maximum nest density in the summer, and again coalesces in the winter, returning to the same winter location year after year. The trails show spatio‐temporal variation as well, depending on the location of nesting and foraging sites. Furthermore, nest movements may be driven by soil microclimate and proximity to man‐made structures. 4. In total, 119 ant nests were discovered in a 3.15 ha plot, 90 (76%) of which belonged to T. sessile. Tapinoma sessile exhibited strong colony connectivity as 78/90 (87%) of nests were connected to at least one other nest by a trail. Mean persistence time for T. sessile nests was 133 ± 5 days. 5. Results indicate that T. sessile is a highly adaptable native ant species that exhibits a high degree of flexibility in its colony social structure. A high degree of polygyny and polydomy may contribute to its ecological dominance and pest status in urban environments.     

Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. The mechanisms by which Argentine ants effectively compete against native ant species have been previously addressed in field studies that centered on interference and exploitation competition at baits and mainly examined the colony-level performance of Argentine ants. Detailed behavioral observations explaining the basis for the strong competitive ability of L. humile are comparatively rare. To gain a better understanding of the mechanisms by which Argentine ants displace native ants we examined the aggressive interactions between the Argentine ants and the odorous house ant, Tapinoma sessile in four different aggression assays: (1) worker dyad interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a resource competition assay which focused on competition for food and nesting space. Our results demonstrate a clear disparity between worker-level and colony-level fighting ability of Argentine ants and provide behavioral evidence to explain the superior interference ability of Argentine ants in group assays. Argentine ants experienced mixed success in fighting against odorous house ants in dyad interactions, but gradually gained a numerical advantage in symmetrical group interactions by active cooperation among nestmates. Results of the resource competition assay indicate that Argentine ants recruit rapidly, numerically dominate food and nesting sites, and aggressively displace T. sessile from baits. Taken together, the results of these assays allow us to pinpoint the behavioral mechanisms responsible for the remarkable competitive ability of Argentine ants.     

The influence of forager number and colony size on food distribution in the odorous house ant, Tapinoma sessile

Stomodeal trophallaxis plays a major role in ant colony nutrition and communication. While the rate of food distribution at the individual level (worker to worker) is rapid, factors affecting the rate of food distribution at the colony level remain poorly understood. We used the odorous house ant, Tapinoma sessile (Say), as a model species to investigate the factors affecting the rate of spread of liquid carbohydrate food throughout a colony. To track the movement of the food we used protein marking and double antibody sandwich enzyme-linked immunosorbent assay, DAS-ELISA. Increasing colony size while keeping the number of donor workers constant significantly decreased the number of individuals testing positive for the marker. After 8 h of trophallactic interactions with ten donors, 92 ± 5% of recipient workers tested positive in a colony of 125 and 38 ± 5% tested positive in a colony of 1,000. Interestingly, as colony size increased and the percentage of workers testing positive decreased, the proportion of workers actually receiving food increased. Food originating from a single donor fed approximately 12 individuals in colonies comprised of 125 recipients and approximately 38 individuals in colonies comprised of 1,000 recipients. Thus, the per capita consumption of food decreased as colony size increased, most likely because the amount of food reaching the colony was limited. Increasing the number of donors while keeping colony size constant significantly increased the number of recipient ants testing positive for the marker. As the number of donor workers doubled, the percentage of recipients testing positive more than doubled suggesting that the number of individuals receiving food increases with increasing colony size, while the per capita amount of food decreases. When food was available ad libitum and in close proximity to the nest, numerous workers fed directly at the food source. This dramatically increased the rate and the extent of food distribution to both the workers and the queens and colony size had no significant effect on the spread of the marker in the workers or the queens. The rate and the extent of food distribution at the colony level may depend on a number of factors including the number of successful foragers, the size and density of the recipient colony, and the recipient caste.     

Dispersed central-place foraging in the polydomous odorous house ant, Tapinoma sessile as revealed by a protein marker

The odorous house ant, Tapinoma sessile, is a native ant species common throughout North America. In urban areas, this ant is classified a pest species and exhibits several attributes characteristic of invasive “tramp” ants (sensu Passera, 1994). These include: extreme polygyny, colony reproduction by budding, reduced internest aggression, generalist diet, and polydomy. Here we explore the organization of foraging and the pathways of food distribution in polydomous colonies of T. sessile in the laboratory and field using a novel marking technique (rabbit IgG protein) and enzyme linked immunosorbent assay (ELISA). Laboratory assays revealed patterns of food allocation from foragers to other castes and developmental stages. Foragers distributed the IgG- labelled sucrose to the majority of workers within 24 h, and workers retained significantly more sucrose than either queens or larvae. Approximately 50% of queens tested positive for the IgG marker and some queens received significantly more sucrose than others, indicating a possible reproductive dominance hierarchy. Larvae received little sucrose demonstrating their minor reliance on carbohydrates. The results of field experiments showed that odorous house ants are dispersed central-place foragers whereby ants from individual nests exhibit high foraging site fidelity, travel along well-established trails, and forage on a local scale. Dispersed central-place foraging most likely allows the odorous house ant to more efficiently secure both clumped and dispersed food sources and possibly increases its competitive ability. As a result, colonies become numerically large and ecologically dominant. The results of our study contribute to our understanding of the social behavior and colony organization in T. sessile. In addition, they provide a framework for designing more effective ant control programs based on liquid baits. Received 13 December 2005; revised 28 February 2006; accepted 3 March 2006.     

Speed-versus-accuracy trade-offs during nest relocation in Argentine ants (Linepithema humile) and odorous house ants (Tapinoma sessile)

Animals are often forced to accommodate disturbance to their territories or nests. When nest relocation becomes necessary, it is important to efficiently evaluate alternative nest sites to choose the one most suitable under current conditions. However, if time is limiting, species may experience a speed-versus-accuracy trade-off when searching for a new home. We examined nest site selection under duress (in the form of flooding) in two species of ants: Linepithema humile and Tapinoma sessile. We predicted that if ants are able to assess and evacuate to the most suitable location, colonies should move to higher elevation, relative to their current nest site, in response to flooding. To test for a speed-versus-accuracy trade-off, we presented colonies with new nest chambers that were either higher, lower, or at the same height as their current nest and examined if their ability to efficiently choose a new site was influenced by the rate of flooding. When flooding rates were slow, both species favored the highest nest site and nearly always moved their entire nest to the same chamber. However, when the rate of flooding was doubled, colonies of T. sessile less often chose the highest nest site and were also more likely to split their nests between two of the available chambers. These results demonstrate a trade-off between speed and accuracy in nest site selection for odorous house ants, while L. humile retained their ability to adequately assess new nest sites under the conditions we presented. These patterns may arise from differences in exploratory behavior and activity between the two species. Despite having identical colony sizes, L. humile had approximately ten times more workers exploring the alternate nest sites 30 min into the experiment than did T. sessile.     

Individual versus social complexity, with particular reference to ant colonies

Insect societies colonies of ants, bees, wasps and termites--vary enormously in their social complexity. Social complexity is a broadly used term that encompasses many individual and colony-level traits and characteristics such as colony size, polymorphism and foraging strategy. A number of earlier studies have considered the relationships among various correlates of social complexity in insect societies; in this review, we build upon those studies by proposing additional correlates and show how all correlates can be integrated in a common explanatory framework. The various correlates are divided among four broad categories (sections). Under 'polyphenism' we consider the differences among individuals, in particular focusing upon 'caste' and specialization of individuals. This is followed by a section on 'totipotency' in which we consider the autonomy and subjugation of individuals. Under this heading we consider various aspects such as intracolony conflict, worker reproductive potential and physiological or morphological restrictions which limit individuals' capacities to perform a range of tasks or functions. A section entitled 'organization of work' considers a variety of aspects, e.g. the ability to tackle group, team or partitioned tasks, foraging strategies and colony reliability and efficiency. A final section, 'communication and functional integration', considers how individual activity is coordinated to produce an integrated and adaptive colony. Within each section we use illustrative examples drawn from the social insect literature (mostly from ants, for which there is the best data) to illustrate concepts or trends and make a number of predictions concerning how a particular trait is expected to correlate with other aspects of social complexity. Within each section we also expand the scope of the arguments to consider these relationships in a much broader sense of'sociality' by drawing parallels with other 'social' entities such as multicellular individuals, which can be understood as 'societies' of cells. The aim is to draw out any parallels and common causal relationships among the correlates. Two themes run through the study. The first is the role of colony size as an important factor affecting social complexity. The second is the complexity of individual workers in relation to the complexity of the colony. Consequently, this is an ideal opportunity to test a previously proposed hypothesis that 'individuals of highly social ant species are less complex than individuals from simple ant species' in light of numerous social correlates. Our findings support this hypothesis. In summary, we conclude that, in general, complex societies are characterized by large colony size, worker polymorphism, strong behavioural specialization and loss of totipotency in its workers, low individual complexity, decentralized colony control and high system redundancy, low individual competence, a high degree of worker cooperation wher tackling tasks, group foraging strategies, high tempo, multi-chambered tailor-made nests, high functional integration, relatively greater use of cues and modulatory signals to coordinate individuals and heterogeneous patterns of worker-worker interaction.     

Responses of ants to disturbance in Australia,with particular reference to functional groups

Abstract Ants are widely used as bioindicators in environmental assessment in Australia, partly because the responses of ant communities to disturbance are relatively well understood. In particular, the use of functional groups has provided a predictive framework for analysing ant community responses to disturbance in the absence of reliable information on the responses of individual species. Here we review 45 studies of the responses of Australian ant communities to disturbance, in order to: (i) identify individual species or species‐groups that respond consistently to disturbance; and (ii) examine the usefulness of the functional group scheme as a framework for predicting ant community responses under different disturbance regimes in different biogeographical regions. The most common forms of disturbance in our studies were fire (17 studies), mining (12; mostly studies of minesite restoration) and grazing (7), with other disturbances including clearing, logging, flooding, recreation, urbanization and farming. Responses of individual species were inevitably variable because of differences in vegetation type, severity of disturbance and time since disturbance. However, we identified a range of widespread species that showed predictable responses, including species of the metallica group of Rhytidoponera in temperate Australia (‘increasers’ in relation to disturbance), species of the terebrans and denticulatus groups of Camponotus (increasers), the aeneovirens group of Melophorus (increasers) from the arid zone, and Iridomyrmex pallidus (increaser) from the monsoonal region. The functional group scheme assessed here was not designed specifically in the context of disturbance, but nevertheless in some situations provides a useful framework for analysing ant community responses. Three distinct syndromes of functional group responses can be recognized. First, Dominant Dolichoderinae and Hot Climate Specialists are groups that prefer open environments, and tend to be favoured by low levels of disturbance in well‐forested habitats. Second, Opportunists and often also Generalized Myrmicinae are broadly adapted taxa with wide habitat tolerances, but are particularly sensitive to competitive interactions such that their responses oppose those of Dominant Dolichoderinae. Finally, Cryptic Species and Specialist Predators have highly specialized requirements that make them especially sensitive to disturbance. Functional groups are of most use in situations where disturbance causes substantial change in habitat structure, particularly in the ground‐layer. Functional groups are of least use in very open habitats, where disturbance merely increases what is already extensive bare ground, and has relatively little impact on microclimate. The ant functional group scheme can play an important role in assessing disturbance in mesic Australia, but may be of more limited use for this in the arid zone.     

Influences on the structure of suburban ant (Hymenoptera: Formicidae) communities and the abundance of Tapinoma sessile

Urbanization can alter the organization of ant communities and affect populations of urban pest ants. In this study, we sampled ant communities in urban and suburban yards to understand the habitat factors that shape these communities and influence the abundance of a common pest species, Tapinoma sessile (Say). We used pitfall traps to sample ant communities and a combination of pitfall traps and baiting to collect T. sessile at 24 sites in Knoxville, TN. In total, we collected 46 ant species. Ant species richness ranged from seven to 24 species per yard. Ant species richness tended to be lowest near houses, whereas T. sessile abundance was highest near houses. The best predictors of ant species richness in yards were canopy cover and presence of leaf litter: ant species richness peaked at mid-levels of canopy cover and was negatively correlated with the presence of leaf litter. Tapinoma sessile abundance increased with presence of logs, boards, or landscaping timbers and leaf litter in yards. Our results indicate that ant communities and the abundance of particular pest species in these urban and suburban landscapes are shaped by many of the same factors that structure ant communities in less anthropogenically disturbed environments.     

Use of terrestrial invertebrates for biodiversity monitoring in Australian rangelands,with particular reference to ants

Abstract Taken literally, the aim of biodiversity monitoring is to track changes in the biological integrity of ecosystems. Given the overwhelmingly dominant contribution of invertebrates to biodiversity, no biodiversity monitoring programme can be considered credible if invertebrates are not addressed effectively. Here we review the use of terrestrial invertebrates, with a particular focus on ants, as bioindicators in Australia in the context of monitoring biodiversity in Australia's rangelands. Ant monitoring systems in Australia were initially developed for assessing restoration success following mining, and have since been applied to a wide range of other land‐use situations, including grazing impacts in rangelands. The use of ants as bioindicators in Australia is supported by an extensive portfolio of studies of the responses of ant communities to disturbance, as well as by a global model of ant community dynamics based on functional groups in relation to environmental stress and disturbance. Available data from mining studies suggest that ants reflect changes in other invertebrate groups, but this remains largely undocumented in rangelands. The feasibility of using ants as indicators in land management remains a key issue, given the large numbers of taxonomically challenging specimens in samples, and a lack of invertebrate expertise within most land‐management agencies. However, recent work has shown that major efficiencies can be achieved by simplifying the ant sorting process, and such efficiencies can actually enhance rather than compromise indicator performance.     

Effects of aromatic cedar mulch on the Argentine ant and the odorous house ant (Hymenoptera: Formicidae).

In laboratory studies, the Argentine ant, Linepithema humile (Mayr), and the odorous house ant, Tapinoma sessile (Say), avoided aromatic cedar mulch as a nesting substrate. Both ant species were killed when confined with fresh aromatic cedar mulch in sealed containers. However, when confined with cedar mulch that had been aged outdoors for up to 140 d, mortality of L. humile was complete regardless of mulch age, whereas T. sessile mortality declined significantly over the mulch-aging period. Argentine ant susceptibility to aromatic cedar mulch was also greater than that of the odorous house ant when colonies were restricted to mulch in open trays. In addition, commercial aromatic cedar oil was lethal to both ant species. Our results suggest that aromatic cedar mulch may serve as an effective component of a comprehensive urban ant management program.     

Recolonization by ants of rehabilitated mineral sand mines on North Stradbroke Island,Queensland, with particular reference to seed removal

The ant fauna of 12 sand-mined plots representing a range of rehabilitation ages and three undisturbed vegetation controls was surveyed during 1982. Physical and botanical parameters were also measured in each plot. Sixty-four ant species were collected from the 15 study plots, of which 44 had colonized one or more of the mined plots. Ant recolonization proceeded rapidly in plots up to 6 years old and may have been influenced by the passage of time, plant cover, density and diversity variables, the amount of litter and by the paucity of logs. In terms of ant species composition, the mined plots were most different from the undisturbed areas. The older plots exhibited a lower ant species richness and this is believed to have resulted from interspecific competition with the tramp ant, Pheidole megacephala. Ant succession proceeded in a slower fashion in the plots dominated by P megacephala. The influence of ants on seeds applied during broadcast seeding of rehabilitated areas was also investigated. Seed removal by ants was greatest for the arillate seeds of Acacia concurrens but moderate quantities of Allocasuarina spp., Eucalyptus spp. Xanthorrhoea sp. and Banksia spp. were taken also. Seed removal by ants was low in the areas which had recently had topsoil applied except where ants foraged from adjacent rehabilitation areas; here they exerted their influence up to 50 m across the fresh topsoil. Seed removal rates in topsoil adjacent to forest were low. Removal rates in a revegetated area 2.5 years old approached those in forest. This indicates that the previous ant-seed relationship had been partially restored by this time, although the relative contribution of seed harvesters and elaiosome collectors still needs to be assessed.     

Biogeography of limno-terrestrial Tardigrada, with particular reference to the Antarctic fauna

The Tardigrada is a cosmopolitan phylum of pre-Pangaean origin, yet tardigrade families and genera show distinct biogeographic components isolated by two major geological events. Separate Laurasian and Gondwanan familial clusters correlate with the Triassic disintegration of Pangaea, while discrete Antarctic, Australian and New Zealand familial/generic clusters relate to the subsequent Jurassic/Cretaceous disintegration of Gondwana.