Critical thermal limits and temperature-dependent walking speed may mediate coexistence between the native winter ant (Prenolepis imparis) and the invasive Argentine ant (Linepithemahumile).

Comparing the thermal tolerance and performance of native and invasive species from varying climatic origins may explain why some native and invasive species can coexist. We compared the thermal niches of an invasive and native ant species. The Argentine ant (Linepithema humile) is an invasive species that has spread to Mediterranean climates worldwide, where it is associated with losses in native arthropod biodiversity. In northern California, long-term surveys of ant biodiversity have shown that the winter ant (Prenolepis imparis) is the native species best able to coexist with Argentine ants. Both species tend hemipteran scales for food, and previous research suggests that these species' coexistence may depend on seasonal partitioning: winter ants are active primarily in the colder winter months, while Argentine ants are active primarily in the warmer months in northern California. We investigated the physiological basis of seasonal partitioning in Argentine and winter ants by a) measuring critical thermal limits, and b) comparing how ant walking speed varies with temperature. While both species had similar CTmax values, we found differences between the two species' critical thermal minima that may allow winter ants to remain functional at ecologically relevant temperatures between 0 and 2.5 °C. We also found that winter ants' walking speeds are significantly less temperature-dependent than those of Argentine ants. Winter ants walk faster than Argentine ants at low temperatures, which may allow the winter ants to remain active and forage at lower winter temperatures. These results suggest that partitioning based on differences in temperature tolerance promotes the winter ant's continued occupation of areas invaded by the Argentine ant.

Foraging behavior and locomotion of the invasive Argentine ant from winter aggregations.

The collective behavior of ant colonies, and locomotion of individuals within a colony, both respond to changing conditions. The invasive Argentine ant (Linepithema humile) thrives in Mediterranean climates with hot, dry summers and colder, wet winters. However, its foraging behavior and locomotion has rarely been studied in the winter. We examined how the foraging behavior of three distinct L. humile colonies was related to environmental conditions and the locomotion of workers during winter in northern California. We found that colonies foraged most between 10 and 15°C, regardless of the maximum daily temperature. Worker walking speed was positively associated with temperature (range 6-24°C) and negatively associated with humidity (range 25-93%RH). All colonies foraged during all day and night hours in a predictable daily cycle, with a correlation between the rate of incoming and outgoing foragers. Foraging activity was unrelated to the activity of a competing native ant species, Prenolepis imparis, which was present in low abundance, and ceased only during heavy rain when ants left foraging trails and aggregated in small sheltered areas on trees.

Distributed nestmate recognition in ants.

We propose a distributed model of nestmate recognition, analogous to the one used by the vertebrate immune system, in which colony response results from the diverse reactions of many ants. The model describes how individual behaviour produces colony response to non-nestmates. No single ant knows the odour identity of the colony. Instead, colony identity is defined collectively by all the ants in the colony. Each ant responds to the odour of other ants by reference to its own unique decision boundary, which is a result of its experience of encounters with other ants. Each ant thus recognizes a particular set of chemical profiles as being those of non-nestmates. This model predicts, as experimental results have shown, that the outcome of behavioural assays is likely to be variable, that it depends on the number of ants tested, that response to non-nestmates changes over time and that it changes in response to the experience of individual ants. A distributed system allows a colony to identify non-nestmates without requiring that all individuals have the same complete information and helps to facilitate the tracking of changes in cuticular hydrocarbon profiles, because only a subset of ants must respond to provide an adequate response.

Water stress strengthens mutualism among ants, trees, and scale insects.

Abiotic environmental variables strongly affect the outcomes of species interactions. For example, mutualistic interactions between species are often stronger when resources are limited. The effect might be indirect: water stress on plants can lead to carbon stress, which could alter carbon-mediated plant mutualisms. In mutualistic ant-plant symbioses, plants host ant colonies that defend them against herbivores. Here we show that the partners' investments in a widespread ant-plant symbiosis increase with water stress across 26 sites along a Mesoamerican precipitation gradient. At lower precipitation levels, Cordia alliodora trees invest more carbon in Azteca ants via phloem-feeding scale insects that provide the ants with sugars, and the ants provide better defense of the carbon-producing leaves. Under water stress, the trees have smaller carbon pools. A model of the carbon trade-offs for the mutualistic partners shows that the observed strategies can arise from the carbon costs of rare but extreme events of herbivory in the rainy season. Thus, water limitation, together with the risk of herbivory, increases the strength of a carbon-based mutualism.

Fast and flexible: argentine ants recruit from nearby trails.

Argentine ants (Linepithema humile) live in groups of nests connected by trails to each other and to stable food sources. In a field study, we investigated whether some ants recruit directly from established, persistent trails to food sources, thus accelerating food collection. Our results indicate that Argentine ants recruit nestmates to food directly from persistent trails, and that the exponential increase in the arrival rate of ants at baits is faster than would be possible if recruited ants traveled from distant nests. Once ants find a new food source, they walk back and forth between the bait and sometimes share food by trophallaxis with nestmates on the trail. Recruiting ants from nearby persistent trails creates a dynamic circuit, like those found in other distributed systems, which facilitates a quick response to changes in available resources.

Diversification and phylogeographic structure in widespread Azteca plant-ants from the northern Neotropics.

The Neotropical myrmecophytic tree Cordia alliodora hosts symbiotic Azteca ants in most of its widespread range. The taxonomy of the genus Azteca is notoriously difficult, which has frequently obscured species identity in ecological studies. We used sequence data from one mitochondrial and four nuclear loci to infer phylogenetic relationships, patterns of geographic distribution, and timing of diversification for 182 colonies of five C. alliodora-dwelling Azteca species from Mexico to Colombia. All morphological species were recovered as monophyletic, but we identified at least five distinct genetic lineages within the most abundant and specialized species, Azteca pittieri. Mitochondrial and nuclear data were concordant at the species level, but not within species. Divergence time analyses estimated that C. alliodora-dwelling Azteca shared a common ancestor approximately 10-22million years ago, prior to the proposed arrival of the host tree in Middle America. Diversification in A. pittieri occurred in the Pleistocene and was not correlated with geographic distance, which suggests limited historical gene flow among geographically restricted populations. This contrasts with the previously reported lack of phylogeographic structure at this spatial scale in the host tree. Climatic niches, and particularly precipitation-related variables, did not overlap between the sites occupied by northern and southern lineages of A. pittieri. Together, these results suggest that restricted gene flow among ant populations may facilitate local adaptation to environmental heterogeneity. Differences in population structure between the ants and their host trees may profoundly affect the evolutionary dynamics of this widespread ant-plant mutualism.

Plant defense, herbivory, and the growth of Cordia alliodora trees and their symbiotic Azteca ant colonies.

The effects of herbivory on plant fitness are integrated over a plant's lifetime, mediated by ontogenetic changes in plant defense, tolerance, and herbivore pressure. In symbiotic ant-plant mutualisms, plants provide nesting space and food for ants, and ants defend plants against herbivores. The benefit to the plant of sustaining the growth of symbiotic ant colonies depends on whether defense by the growing ant colony outpaces the plant's growth in defendable area and associated herbivore pressure. These relationships were investigated in the symbiotic mutualism between Cordia alliodora trees and Azteca pittieri ants in a Mexican tropical dry forest. As ant colonies grew, worker production remained constant relative to ant-colony size. As trees grew, leaf production increased relative to tree size. Moreover, larger trees hosted lower densities of ants, suggesting that ant-colony growth did not keep pace with tree growth. On leaves with ants experimentally excluded, herbivory per unit leaf area increased exponentially with tree size, indicating that larger trees experienced higher herbivore pressure per leaf area than smaller trees. Even with ant defense, herbivory increased with tree size. Therefore, although larger trees had larger ant colonies, ant density was lower in larger trees, and the ant colonies did not provide sufficient defense to compensate for the higher herbivore pressure in larger trees. These results suggest that in this system the tree can decrease herbivory by promoting ant-colony growth, i.e., sustaining space and food investment in ants, as long as the tree continues to grow.

Effects of vegetation cover, presence of a native ant species, and human disturbance on colonization by Argentine ants.

The spread of non-native invasive species is affected by human activity, vegetation cover, weather, and interaction with native species. We analyzed data from a 17-year study of the distribution of the non-native Argentine ant (Linepithema humile) and the native winter ant (Prenolepis imparis) in a preserve in northern California (U.S.A.). We conducted logistic regressions and used model selection to determine whether the following variables were associated with changes in the distribution of each species: presence of conspecifics at neighboring sites, distance to development (e.g., roads, buildings, and landscaped areas), proportion of vegetation cover taller than 0.75 m, elevation, distance to water, presence of both species at a site, temperature, and rainfall. Argentine ants colonized unoccupied sites from neighboring sites, but the probability of appearance and persistence decreased as distance to development, vegetation cover, and elevation increased. Winter ants appeared and persisted in sites with relatively high vegetation cover (i.e., highly shaded sites). Presence of the 2 species was negatively associated in sites with high vegetation cover (more winter ants) and sites near development (more Argentine ants). Probability of colonization of Argentine ants decreased where winter ants were most persistent. At sites near development within the preserve, abundant Argentine ant populations may be excluding winter ants. The high abundance of Argentine ants at these sites may be due to immigration from suburban areas outside the preserve, which are high-quality habitat for Argentine ants. In the interior of the preserve, distance from development, low-quality habitat, and interaction with winter ants may in combination exclude Argentine ants. Interactions among the variables we examined were associated with low probabilities of Argentine ant colonization in the preserve.

Chemical defense by the native winter ant (Prenolepis imparis) against the invasive Argentine ant (Linepithema humile).

The invasive Argentine ant (Linepithema humile) is established worldwide and displaces native ant species. In northern California, however, the native winter ant (Prenolepis imparis) persists in invaded areas. We found that in aggressive interactions between the two species, P. imparis employs a potent defensive secretion. Field observations were conducted at P. imparis nest sites both in the presence and absence of L. humile. These observations suggested and laboratory assays confirmed that P. imparis workers are more likely to secrete when outnumbered by L. humile. Workers of P. imparis were also more likely to secrete near their nest entrances than when foraging on trees. One-on-one laboratory trials showed that the P. imparis secretion is highly lethal to L. humile, causing 79% mortality. The nonpolar fraction of the secretion was chemically analyzed with gas chromatography/mass spectrometry, and found to be composed of long-chain and cyclic hydrocarbons. Chemical analysis of dissected P. imparis workers showed that the nonpolar fraction is derived from the Dufour's gland. Based on these conclusions, we hypothesize that this chemical defense may help P. imparis to resist displacement by L. humile.

Rainfall facilitates the spread, and time alters the impact, of the invasive Argentine ant.

Climate change may exacerbate invasions by making conditions more favorable to introduced species relative to native species. Here we used data obtained during a long-term biannual survey of the distribution of ant species in a 481-ha preserve in northern California to assess the influence of interannual variation in rainfall on the spread of invasive Argentine ants, Linepithema humile, and the displacement of native ant species. Since the survey began in 1993, Argentine ants have expanded their range into 74 new hectares. Many invaded hectares were later abandoned, so the range of Argentine ants increased in some years and decreased in others. Rainfall predicted both range expansion and interannual changes in the distribution of Argentine ants: high rainfall, particularly in summer months, promoted their spread in the summer. This suggests that an increase in rainfall will promote a wider distribution of Argentine ants and increase their spread into new areas in California. Surprisingly, the distribution of two native ant species also increased following high rainfall, but only in areas of the preserve that were invaded by L. humile. Rainfall did not have a negative impact on total native ant species richness in invaded areas. Instead, native ant species richness in invaded areas increased significantly over the 13 years of observation. This suggests that the impact of Argentine ants on naïve ant communities may be most severe early in the invasion process.

The devil to pay: a cost of mutualism with Myrmelachista schumanni ants in 'devil's gardens' is increased herbivory on Duroia hirsuta trees.

'Devil's gardens' are nearly pure stands of the myrmecophyte, Duroia hirsuta, that occur in Amazonian rainforests. Devil's gardens are created by Myrmelachista schumanni ants, which nest in D. hirsuta trees and kill other plants using formic acid as an herbicide. Here, we show that this ant-plant mutualism has an associated cost; by making devil's gardens, M. schumanni increases herbivory on D. hirsuta. We measured standing leaf herbivory on D. hirsuta trees and found that they sustain higher herbivory inside than outside devil's gardens. We also measured the rate of herbivory on nursery-grown D. hirsuta saplings planted inside and outside devil's gardens in ant-exclusion and control treatments. We found that when we excluded ants, herbivory on D. hirsuta was higher inside than outside devil's gardens. These results suggest that devil's gardens are a concentrated resource for herbivores. Myrmelachista schumanni workers defend D. hirsuta against herbivores, but do not fully counterbalance the high herbivore pressure in devil's gardens. We suggest that high herbivory may limit the spread of devil's gardens, possibly explaining why devil's gardens do not overrun Amazonian rainforests.

Ecology: 'Devil's gardens' bedevilled by ants.

'Devil's gardens' are large stands of trees in the Amazonian rainforest that consist almost entirely of a single species, Duroia hirsuta, and, according to local legend, are cultivated by an evil forest spirit. Here we show that the ant Myrmelachista schumanni, which nests in D. hirsuta stems, creates devil's gardens by poisoning all plants except its host plants with formic acid. By killing these other plants, M. schumanni provides its colonies with abundant nest sites--a long-lasting benefit as colonies can live for 800 years.

Long-term dynamics of the distribution of the invasive Argentine ant, Linepithema humile, and native ant taxa in northern California.

Invasive species, where successful, can devastate native communities. We studied the dynamics of the invasive Argentine ant, Linepithema humile, for 7 years in Jasper Ridge, a biological preserve in northern California. We monitored the distributions at the hectare scale of native ant taxa and L. humile in the spring and fall from 1993 to 1999. We also studied the invasion dynamics at a finer resolution by searching for ants in 1-m2 plots. Our results are similar at both scales. The distributions of several native species are not random with regard to L. humile; the distributions of several epigeic species with similar habitat affinities overlap much less frequently than expected with the distribution of L. humile. We found that season had a significant influence on the distributions of L. humile and several native taxa. Over the 7-year period, L. humile has increased its range size in Jasper Ridge largely at the expense of native taxa, but there is seasonal and yearly variation in this rate of increase. Studies of invasions in progress which sample across seasons and years may help to predict the spread and effects of invasive species.

Exploitation and interference competition between the invasive Argentine ant, Linepithema humile, and native ant species.

Interactions between the invasive Argentine ant, Linepithema humile, and native ant species were studied in a 450-ha biological reserve in northern California. Along the edges of the invasion, the presence of Argentine ants significantly reduced the foraging success of native ant species, and vice versa. Argentine ants were consistently better than native ants at exploiting food sources: Argentine ants found and recruited to bait more consistently and in higher numbers than native ant species, and they foraged for longer periods throughout the day. Native ants and Argentine ants frequently fought when they recruited to the same bait, and native ant species were displaced from bait during 60% of these encounters. In introduction experiments, Argentine ants interfered with the foraging of native ant species, and prevented the establishment of new colonies of native ant species by preying upon winged native ant queens. The Argentine ants' range within the preserve expanded by 12 ha between May 1993 and May 1994, and 13 between September 1993 and September 1994, with a corresponding reduction of the range of native ant species. Although some native ants persist locally at the edges of the invasion of Argentine ants, most eventually disappear from invaded areas. Both interference and exploitation competition appear to be important in the displacement of native ant species from areas invaded by Argentine ants.