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Natural terrain is rarely flat. Substrate irregularities challenge walking animals to maintain stability, yet we lack quantitative assessments of walking performance and limb kinematics on naturally uneven ground. We measured how continually uneven 3D-printed substrates influence walking performance of Argentine ants by measuring walking speeds of workers from laboratory colonies and by testing colony-wide substrate preference in field experiments. Tracking limb motion in over 8000 videos, we used statistical models that associate walking speed with limb kinematic parameters to compare movement over flat versus uneven ground of controlled dimensions. We found that uneven substrates reduced preferred and peak walking speeds by up to 42% and that ants actively avoided uneven terrain in the field. Observed speed reductions were modulated primarily by shifts in stride frequency instead of stride length (flat R 2: 0.91 versus 0.50), a pattern consistent across flat and uneven substrates. Mixed effect modelling revealed that walking speeds on uneven substrates were accurately predicted based on flat walking data for over 89% of strides. Those strides that were not well modelled primarily involved limb perturbations, including missteps, active foot repositioning and slipping. Together these findings relate kinematic mechanisms underlying walking performance on uneven terrain to ecologically relevant measures under field conditions.

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Flooding impacts ground nesting ant colonies by destroying the infrastructure housing and organizing societal function. Here, we report the convergent evolution in distantly related ant species of a behavioral trait that minimizes costs of flooding: the construction of earthen levees around nest entrances. In a South American floodplain ecosystem, we observed five ant species constructing prominent earthen berms encircling nest entrances shortly after large rainfall events. In four of these species, experimental flooding of nests demonstrated that earthen berms sufficed to prevent floodwaters from entering the below ground portions of the nest. Additional manipulations revealed that levee breaching caused, pronounced, and extended reductions in food collection for two distantly related species. Foraging was preempted by the allocation of workers to repair the internal structure of the nest. These findings represent convergent evolution of a functionally important nest construction behavior in response to comparable selective forces.

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Variation in invasion success may result from the divergent evolutionary histories of introduced species compared to those of native taxa. The vulnerability of native biotas to ecological disruption may be especially great on oceanic islands invaded by continental species with unique ecological traits. In part because Hawaii lacks native eusocial insects, social invaders may threaten endemic taxa that are ecologically similar but solitary. Using a combination of field manipulations, molecular analyses, physiological data, and behavioral assays, we identify the mechanisms underlying the displacement of two genera of native solitary Hymenoptera in Hawaii by a social continental invader, the western yellowjacket (Vespula pensylvanica). Experimental removal of V. pensylvanica colonies resulted in increased densities of native Hymenoptera. Endemic Hylaeus bees directly suffer through predation by yellowjackets, and perhaps as a consequence, avoid floral resources occupied by V. pensylvanica. Native Nesodynerus wasps also avoid V. pensylvanica but are negatively affected by yellowjackets not through predation, but through exploitative competition for caterpillar prey. Displacement of native solitary Hymenoptera may be heightened by the ability of V. pensylvanica to prey upon and scavenge honey bees and to rob their honey stores, resources unavailable to endemic bees and wasps because of their specialized niches. Our study provides a unique example of an ecologically generalized social invader that restructures native assemblages of solitary Hymenoptera by interacting with endemic taxa on multiple trophic levels.

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Baker and Stebbins's 1965 book The Genetics of Colonizing Species aimed to draw together scientists from a variety of disciplines to provide a conceptual framework for the study of species introductions. A goal of their volume was to examine how studies on biological invasions could be used to provide insight into basic research questions as well as to develop practical strategies for control. In this article, we attempt to follow the goals of Baker and Stebbins by reviewing work on the genetics and behavior of a widespread colonizing species, the Argentine ant (Linepithema humile). Specifically, we examine the evolutionary changes that have taken place as a result of this species being introduced into new environments and synthesize recent research on Argentine ants from the perspective of population genetics, recognition systems, and the mechanisms that may underlie their ecological success.

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Social insects can modulate body temperature to increase foraging efficiency; however, little is known about how the relative value of protein resources affects forager body temperature. Such regulation may be important given that colony growth is often limited by protein availability. In this paper, we present what are, to our knowledge, the first data for social insects showing that thoracic temperatures (T (th)) of foragers increase with the protein content of food resources. In an introduced population of western yellowjacket (Vespula pensylvanica), we measured T (th) of foragers collecting high-quality protein (100% canned chicken) and low-quality protein (50% canned chicken, 50% indigestible alpha-cellulose by volume) at different ambient air temperatures (T (a)). Wasps foraging on 100% chicken consistently exhibited higher T (th) compared to wasps foraging on 50% chicken. After correcting for T (a), the mean T (th) for wasps collecting 100% chicken were 1.98 degrees C higher than those of individuals collecting 50% chicken. We suggest that this mechanism may increase foraging efficiency in this and other social wasp species.

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An understanding of why introduced species achieve ecological success in novel environments often requires information about the factors that limit the abundance of these taxa in their native ranges. Although numerous recent studies have evaluated the importance of natural enemies in this context, relatively few have examined how ecological success may result from differences in the magnitude of interference competition between communities in the native and introduced ranges of nonnative species. Here we examine how native-range competitive environments may relate to invasion success for two important invasive species, the red imported fire ant (Solenopsis invicta) and the Argentine ant (Linepithema humile), in a region of native-range sympatry. At two study sites in northern Argentina, we used stable-isotope analysis, a variety of observational approaches, and two different reciprocal removal experiments to test (1) whether S. invicta competes asymmetrically with L. humile (as suggested by the 20th century pattern of replacement in the southeastern United States) and (2) the extent to which these two species achieve behavioral and numerical dominance. Stable-isotope analysis and activity surveys indicated that S. invicta and L. humile are both omnivores and forage during broadly overlapping portions of the diel cycle. Short-term removal experiments at baits revealed no competitive asymmetry between S. invicta and L. humile. Longer-term colony removal experiments illustrated that S. invicta and L. humile experience an approximately equal competitive release upon removal of the other. Our results indicate that neither S. invicta nor L. humile achieves the same degree of behavioral or ecological dominance where they co-occur in native populations as they do in areas where either is common in their introduced range. These results strongly suggest that interspecific competition is an important limiting factor for both S. invicta and L. humile in South America.

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Although the ecological success of introduced species hinges on biotic interactions and physical conditions, few experimental studies--especially on animals--have simultaneously investigated the relative importance of both types of factors. The lack of such research may stem from the common assumption that native and introduced species exhibit similar environmental tolerances. Here we combine experimental and spatial modeling approaches (1) to determine the relative importance of biotic and abiotic controls of Argentine ant (Linepithema humile) invasion success, (2) to examine how the importance of these factors changes with spatial scale in southern California (USA), and (3) to assess how Argentine ants differ from native ants in their environmental tolerances. A factorial field experiment that combined native ant removal with irrigation revealed that Argentine ants failed to invade any dry plots (even those lacking native ants) but readily invaded all moist plots. Native ants slowed the spread of Argentine ants into irrigated plots but did not prevent invasion. In areas without Argentine ants, native ant species showed variable responses to irrigation. At the landscape scale, Argentine ant occurrence was positively correlated with minimum winter temperature (but not precipitation), whereas native ant diversity increased with precipitation and was negatively correlated with minimum winter temperature. These results are of interest for several reasons. First, they demonstrate that fine-scale differences in the physical environment can eclipse biotic resistance from native competitors in determining community susceptibility to invasion. Second, our results illustrate surprising complexities with respect to how the abiotic factors limiting invasion can change with spatial scale, and third, how native and invasive species can differ in their responses to the physical environment. Idiosyncratic and scale-dependent processes complicate attempts to forecast where introduced species will occur and how their range limits may shift as a result of climate change.

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The Argentine ant (Linepithema humile) is a damaging invasive species that has become established in many Mediterranean-type ecosystems worldwide. To identify likely sources of introduced populations we examined the relationships among native Linepithema populations from Argentina and Brazil and introduced populations of L. humile using mitochondrial cytochrome b sequence data and nuclear microsatellite allele frequencies. The mitochondrial phylogeny revealed that the populations in Brazil were only distantly related to both the introduced populations and the native populations in Argentina, and confirmed that populations in Brazil, previously identified as L. humile, are likely a different species. The microsatellite-based analysis provided resolution among native and introduced populations of L. humile that could not be resolved using the mitochondrial sequences. In the native range, colonies that were geographically close to one another tended to be genetically similar, whereas more distant colonies were genetically different. Most samples from the introduced range were genetically similar, although some exceptions were noted. Most introduced populations were similar to native populations from the southern Rio Parana and were particularly similar to a population from Rosario, Argentina. These findings implicate populations from the southern Rio Parana as the most likely source of introduced populations. Moreover, these data suggest that current efforts to identify natural enemies of the Argentine ant for biological control should focus on native populations in the southern Rio Parana watershed.

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Invading organisms may spread through local movements (giving rise to a diffusion-like process) and by long-distance jumps, which are often human-mediated. The local spread of invading organisms has been fit with varying success to models that couple local population growth with diffusive spread, but to date no quantitative estimates exist for the relative importance of local dispersal relative to human-mediated long-distance jumps. Using a combination of literature review, museum records, and personal surveys, we reconstruct the invasion history of the Argentine ant (Linepithema humile), a widespread invasive species, at three spatial scales. Although the inherent dispersal abilities of Argentine ants are limited, in the last century, human-mediated dispersal has resulted in the establishment of this species on six continents and on many oceanic islands. Human-mediated jump dispersal has also been the primary mode of spread at a continental scale within the United States. The spread of the Argentine ant involves two discrete modes. Maximum distances spread by colonies undergoing budding reproduction averaged 150 m/year, whereas annual jump-dispersal distances averaged three orders of magnitude higher. Invasions that involve multiple dispersal processes, such as those documented here, are undoubtedly common. Detailed data on invasion dynamics are necessary to improve the predictive power of future modeling efforts.

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Despite the severe ecological and economic damage caused by introduced species, factors that allow invaders to become successful often remain elusive. Of invasive taxa, ants are among the most widespread and harmful. Highly invasive ants are often unicolonial, forming supercolonies in which workers and queens mix freely among physically separate nests. By reducing costs associated with territoriality, unicolonial species can attain high worker densities, allowing them to achieve interspecific dominance. Here we examine the behavior and population genetics of the invasive Argentine ant (Linepithema humile) in its native and introduced ranges, and we provide a mechanism to explain its success as an invader. Using microsatellite markers, we show that a population bottleneck has reduced the genetic diversity of introduced populations. This loss is associated with reduced intraspecific aggression among spatially separate nests, and leads to the formation of interspecifically dominant supercolonies. In contrast, native populations are more genetically variable and exhibit pronounced intraspecific aggression. Although reductions in genetic diversity are generally considered detrimental, these findings provide an example of how a genetic bottleneck can lead to widespread ecological success. In addition, these results provide insights into the origin and evolution of unicoloniality, which is often considered a challenge to kin selection theory.

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