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SignificanceInvasive social insects are among the most damaging of invasive organisms and have proved universally intractable to biological control. Despite this, populations of some invasive social insects collapse from unknown causes. We report long-term studies demonstrating that infection by a microsporidian pathogen causes populations of a globally significant invasive ant to collapse to local extinction, providing a mechanistic understanding of a pervasive phenomenon in biological invasions: the collapse of established populations from endogenous factors. We apply this knowledge and successfully eliminate two large, introduced populations of these ants. More broadly, microsporidian pathogens should be evaluated for control of other supercolonial invasive social insects. Diagnosing the cause of unanticipated population collapse in invasive organisms can lead to applied solutions.

The microsporidian pathogen Myrmecomorba nylanderiae Plowes et al. infects introduced tawny crazy ants (Nylanderia fulva (Mayr)), and constitutes one of the first natural enemies known to attack this invasive ant. We assess how infection is transmitted within colonies and how infection impacts N. fulva colony fragment growth under carbohydrate-deficient and carbohydrate-sufficient dietary conditions. Carbohydrate scarcity is a common source of stress for ant colonies. Infected workers efficiently pass infection to developing larvae but all other potential pathways for within colony transmission are rare or non-viable. For unknown reasons, queens within infected colony fragments are generally uninfected, limiting the role of transovarial transmission in intracolony transmission. In the laboratory, infection by M. nylanderiae primarily impacts the growth of N. fulva colonies by reducing pupal production. Colony growth showed a substantially greater impact under carbohydrate-deficient conditions implying that the effect of the pathogen may depend on seasonally variable carbohydrate availability. In the colony growth assay, worker mortality did not differ with infection status under either nutrient regime. However, in a longer, direct test of survivorship, infected worker survivorship was significantly lower. Recently, some established N. fulva populations with high prevalence of M. nylanderiae infection have declined precipitously, though it is unknown if M. nylanderiae is a causative agent in these declines. The combination of chronic impacts, presence in North America, and potential association with population declines makes M. nylanderiae a promising prospect for the biological control of N. fulva.

How workers within an ant colony perceive and enforce colony boundaries is a defining biological feature of an ant species. Ants fall along a spectrum of social organizations ranging from single-queen, single nest societies to species with multi-queen societies in which workers exhibit colony-specific, altruistic behaviors towards non-nestmate workers from distant locations. Defining where an ant species falls along this spectrum is critical for understanding its basic ecology. Herein we quantify queen numbers, describe intraspecific aggression, and characterize the distribution of colony sizes for tawny crazy ant (Nylanderia fulva) populations in native range areas in South America as well as in their introduced range in the Southeastern United States. In both ranges, multi-queen nests are common. In the introduced range, aggressive behaviors are absent at all spatial scales tested, indicating that within the population in the Southeastern United States N. fulva is unicolonial. However, this contrasts strongly with intraspecific aggression in its South American native range. In the native range, intraspecific aggression between ants from different nests is common and ritualized. Aggression is typically one-sided and follows a stereotyped sequence of escalating behaviors that stops before actual fighting occurs. Spatial patterns of non-aggressive nest aggregation and the transitivity of non-aggressive interactions demonstrate that results of neutral arena assays usefully delineate colony boundaries. In the native range, both the spatial extent of colonies and the average number of queens encountered per nest differ between sites. This intercontinental comparison presents the first description of intraspecific aggressive behavior for this invasive ant and characterizes the variation in colony organization in the native-range, a pre-requisite to a full understanding of the origins of unicoloniality in its introduced range.

Analysis of an invasive species' niche shift between native and introduced ranges, along with potential distribution maps, can provide valuable information about its invasive potential. The tawny crazy ant, Nylanderia fulva, is a rapidly emerging and economically important invasive species in the southern United States. It is originally from east‐central South America and has also invaded Colombia and the Caribbean Islands. Our objectives were to generate a global potential distribution map for N. fulva, identify important climatic drivers associated with its current distribution, and test whether N. fulva's realized climatic niche has shifted across its invasive range. We used MaxEnt niche model to map the potential distribution of N. fulva using its native and invaded range occurrences and climatic variables. We used principal component analysis methods for investigating potential shifts in the realized climatic niche of N. fulva during invasion. We found strong evidence for a shift in the realized climatic niche of N. fulva across its invasive range. Our models predicted potentially suitable habitat for N. fulva in the United States and other parts of the world. Our analyses suggest that the majority of observed occurrences of N. fulva in the United States represent stabilizing populations. Mean diurnal range in temperature, degree days at ≥10°C, and precipitation of driest quarter were the most important variables associated with N. fulva distribution. The climatic niche expansion demonstrated in our study may suggest significant plasticity in the ability of N. fulva to survive in areas with diverse temperature ranges shown by its tolerance for environmental conditions in the southern United States, Caribbean Islands, and Colombia. The risk maps produced in this study can be useful in preventing N. fulva's future spread, and in managing and monitoring currently infested areas.

A recently introduced, ecologically dominant, exotic ant species, Nylanderia fulva, is invading the Southeastern United States and Texas. We evaluate how this invader impacts diversity and abundance of co-occurring ants and other arthropods in two grasslands. N. fulva rapidly attains densities up to 2 orders of magnitude greater than the combined abundance of all other ants. Overall ant biomass increases in invaded habitat, indicating that N. fulva exploits resources not fully utilized by the local ant assemblage. At high density, as N. fulva spreads, it eliminates the current ecologically dominant invasive ant, red imported fire ants (Solenopsis invicta). Compared to imported fire ant dominated habitat, N. fulva invasion zones have lower non-ant arthropod species richness and abundance with impacts differing by trophic category. Further, N. fulva reduces abundance and species richness of the remainder of the ant assemblage and does so in a non-random manner: impacting species with small sized workers much less than species with larger workers. In these and other ant assemblages with a large exotic component, the exotics tend to be small bodied species. As a result, N. fulva almost completely eliminates regionally distributed species, but leaves globally distributed species largely unaffected, thereby systematically favoring introduced over native diversity. S. invicta impacts wildlife and arthropod assemblage structure and is nearly ubiquitous in non-forested habitats of the Southeastern United States and Texas. Its displacement by N. fulva has critical implications for the natural systems of this region.

The functions and compositions of symbiotic bacterial communities often correlate with host ecology. Yet cause-and-effect relationships, and the order of symbiont versus host change, remain unclear in the face of ancient symbioses and conserved host ecology. Several groups of ants exemplify this challenge, as their low-nitrogen diets and specialized symbioses appear conserved and ancient. To address whether nitrogen-provisioning symbionts might be important in the early stages of ant trophic shifts we studied bacteria from the Argentine ant, Linepithema humile—an invasive species that has transitioned toward greater consumption of sugar-rich, nitrogen-poor foods in parts of its introduced range. Bacteria were present at low densities in most L. humile workers, and among those yielding quality 16S rRNA amplicon sequencing data we found just three symbionts to be common and dominant. Two, a Lactobacillus and an Acetobacteriaceae species, were shared between native and introduced populations. The other, a Rickettsia, was found only in two introduced supercolonies. Across an eight-year period of trophic reduction in one introduced population, we found no change in symbionts, arguing against a relationship between natural dietary change and microbiome composition. Overall our findings thus argue against major changes in symbiotic bacteria in association with the invasion and trophic shift of L. humile. In addition, genome content from close relatives of the identified symbionts suggest that just one can synthesize most essential amino acids; this bacterium was only modestly abundant in introduced populations, providing little support for a major role of nitrogen-provisioning symbioses in Argentine ant's dietary shift. This article is protected by copyright. All rights reserved.

Summary 1. Habitat disturbance and species invasions interact in natural systems, making it difficult to isolate the primary cause of ecosystem degradation. A general understanding requires case studies of how disturbance and invasion interact across a variety of ecosystem – invasive species combinations. 2. Dramatic losses in ant diversity followed the invasion of central Texas by red imported fire ants (Solenopsis invicta). However, recent manipulative studies in Florida revealed no effect on ant diversity following the removal of S. invicta from a disturbed pasture habitat, but moderate loss of diversity associated with their introduction into undisturbed habitat and no invasion occurred without disturbance. Thus, the importance of S. invicta in driving diversity loss and its ability to invade undisturbed systems is unresolved. 3. We examine the distribution and abundance of a large monogyne S. invicta population and its association with the co-occurring ant assemblage at a site in south Texas close to the aridity tolerance limit of S. invicta. 4. We document that moisture modulates S. invicta densities. Further, soil disturbing habitat manipulations greatly increase S. invicta population densities. However, S. invicta penetrates all habitats regardless of soil disturbance history. In contrast, controlled burns depress S. invicta densities. 5. In habitats where S. invicta is prevalent, it completely replaces native fire ants. However, S. invicta impacts native ants as a whole less strongly. Intriguingly, native ants responded distinctly to S. invicta in different environments. In wet, undisturbed environments, high S. invicta abundance disrupts the spatial structure of the ant assemblage by increasing clumping and is associated with reduced species density, while in dry-disturbed habitats, sites with high S. invicta abundance possess high numbers of native species. Analyses of co-occurrence indicate that reduced species density in wet-undisturbed sites arises from negative species interactions between native ants and S. invicta. However, these same data suggest that the high native species density of abundant S. invicta sites in dry-disturbed environments does not result from facilitation. 6. Monogyne S. invicta populations play different roles in different environments, driving ant diversity loss in some, but being largely symptomatic of habitat disturbance in others.

Mutualisms play key roles in the functioning of ecosystems. However, reciprocally beneficial interactions that involve introduced species also can enhance invasion success and in doing so compromise ecosystem integrity. For example, the growth and competitive ability of introduced plant species can increase when fungal or microbial associates provide limiting nutrients. Mutualisms also may aid animal invasions, but how such systems may promote invasion success has received relatively little attention. Here we examine how access to food-for-protection mutualisms involving the red imported fire ant ( Solenopsis invicta ) aids the success of this prominent invader. Intense interspecific competition in its native Argentina constrained the ability of S. invicta to benefit from honeydew-producing Hemiptera (and other accessible sources of carbohydrates), whereas S. invicta dominated these resources in its introduced range in the United States. Consistent with this strong pattern, nitrogen isotopic data revealed that fire ants from populations in the United States occupy a lower trophic position than fire ants from Argentina. Laboratory and field experiments demonstrated that honeydew elevated colony growth, a crucial determinant of competitive performance, even when insect prey were not limiting. Carbohydrates, obtained largely through mutualistic partnerships with other organisms, thus represent critical resources that may aid the success of this widespread invasive species. These results illustrate the potential for mutualistic interactions to play a fundamental role in the establishment and spread of animal invasions.

The introduction and establishment of Pseudacteon obtusus Borgmeier (Diptera: Phoridae) into the United States is an important milestone in the effort to introduce a suite of parasitoids and pathogens as prospective biocontrol agents of the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae).P. obtusus is the fourth species of phorid fly to be established in the US and is now present in at least 11 sites in Texas. This species has a greater propensity for attacking foraging worker ants, and to a lesser degree it will attend mound disturbance events. The effectiveness of alternate field release strategies was evaluated across 28 sites in Texas, with establishment success 35 times more likely when no other phorid species were present at the outset of the releases. This result implies that competition between phorid species may interfere with establishment. Therefore release strategies should be directed to locations lacking other naturalized species or to times when abundances of other species are low.

Kneallhazia solenopsae is a pathogenic microsporidium that infects the fire ants Solenopsis invicta and Solenopsis richteri in South America and the USA. In this study, we analyzed the prevalence and molecular diversity of K. solenopsae in fire ants from North and South America. We report the first empirical evidence of K. solenopsae infections in the tropical fire ant, Solenopsis geminata, and S. geminata × Solenopsis xyloni hybrids, revealing an expanded host range for this microsporidium. We also analyzed the molecular diversity at the 16S ribosomal RNA gene in K. solenopsae from the ant hosts S. invicta, S. richteri, S. geminata and S. geminata × S. xyloni hybrids from North America, Argentina and Brazil. We found 22 16S haplotypes. One of these haplotypes (WD_1) appears to be widely distributed, and is found in S. invicta from the USA and S. geminata from southern Mexico. Phylogenetic analyses of 16S sequences revealed that K. solenopsae haplotypes fall into one of two major clades that are differentiated by 2–3%. In some cases, multiple K. solenopsae haplotypes per colony were found, suggesting either an incomplete homogenization among gene copies within the 16S gene cluster or multiple K. solenopsae variants simultaneously infecting host colonies.

Some phorid flies of the genus Pseudacteon Coquillett parasitize workers of Solenopsis geminata (F.) complex fire ants. The group is found to comprise at least 21 species of flies, of which 11 are new to science and described here: P. amuletum, P. andinus, P. annulus, P. catarinae, P. deltoides, P. hippeus, P. kungae, P. laticarinatus, P. palomita, P. quinni, and P. robustus. An identification key to the females is included. This expansion of known Pseudacteon species parasitizing the S. geminata complex makes it an interesting system for comparison with Pseudacteon flies that parasitize fire ants of the S. saevissima (Smith) complex.

The once-extensive tallgrass prairie community of North America has been reduced to small rem- nants, many of which are surrounded by intensive corn (Zea mays) agriculture. We investigated adult corn- rootworm beetles (Chrysomelidae: Diabrotica spp.), important pests of corn, on sunflowers (Asteraceae: He- lianthus spp.) in prairie remnants in southeast Minnesota. Large numbers of beetles invaded the prairie from surrounding corn fields in late summer. D. barberi and D. virgifera were captured on sticky traps in all loca- tions in the prairie, but abundance was much greater near the edge adjacent to corn. We observed D, barberi (but not D. virgifera) feeding extensively on sunflower pollen and occasionally on other flower parts, such as petak. Sunflowers located nearer corn fields sustained morefloral damage than those farther from om. To de- termine the effect of beetle &mage on seed set, we enclosed sunflower heads in bags with either zero, two, or four D. barberi adults. Seed set was reduced in heads enclosed with D. barberi Thus, this agriculturalpest may interfere with the successful reproduction of sunflowers and possibly otherprairie composites thatflower in late summer. Given the small size of mostprairie remnants and the abundance of thisflowerrfeeding beetle in land- scapes dominated by corn agriculture, D. barberi may affect the sustainability ofprairie plantpopulations.

Since the first report on introductions to Texas of Pseudacteon decapitating flies, a variety of participants have released flies in a range of sites. The expansions of Pseudacteon populations have been systematically and widely monitored. Before 2002, the widely released initial species P. tricuspis Borgmeier did not become established. Severe drought in 1996–2001 and host-size-dependent sex ratio were proposed constraints in establishing this species. In recent years, however, these limitations have been lifted in some areas by favorable weather, irrigation of release sites, and/or by use of a smaller Pseudacteon species, P. curvatus Borgmeier, not reliant on larger fire ant workers to produce females. Beginning in 2002, the USDA-APHIS collaboration with USDA-ARS and Texas Cooperative Extension programs began to supplement release sites in Texas beyond those initiated by the University of Texas, Austin phorid fly project. In 2005, private citizens began to participate in the spread of Pseudacte...

Combating invasive species requires a detailed, mechanistic understanding of the manner and speed with which organisms expand their ranges. Biological control efforts provide an opportunity to study the process of species invasions and range expansions under known initial conditions. This study examines the rate, pattern and mechanisms of spread for two populations of the biological control agent Pseudacteon tricuspis, phorid-fly parasitoids of imported fire ants. We employ a trap-based survey method that detects phorid flies in low-density populations, and provides data on abundance. This technique allows us to differentiate between continuous population spread and effective long-distance dispersal and to examine density gradients of phorid flies across the expanding population front. We find that occupied sites in front of the leading edge of continuous populations were common; forming small populations we refer to as satellite populations. Satellite populations are tens of kilometers from the nearest possible source. Wind governs the dynamics of spread in these two central Texas populations. Population edges expanding with the wind exhibited a higher frequency of effective long-distance dispersal than did populations expanding into the wind. This enhanced effective long-distance dispersal rate translated into a five times faster rate of spread for population edges traveling with the wind. This planned invasion shares many characteristics in common with unplanned species invasions including: protracted establishment phase during which densities were below detection thresholds, and slow initial spread immediately after establishment followed by rapid, accelerating spread rates as population sizes grew.

Summary 1 Trade-offs underpin local species coexistence. Trade-offs between interference and exploitative competitive ability provie a mechanism for explaining species coexistence within guilds that exploit overlapping resources. 2 Omnivorous, leaf litter ants exploit a shared food base and occur in species-rich assemblages. In these assemblages, species that excel at usurping food items from other species are poor at finding food items first. In assemblages where some members are attacked by phorid fly parasitoids, host species face an additional trade-off between defending themselves against parasitic attack and maximizing their competitive abilities. Host species thus face two trade-offs that interact via the trait-mediated indirect interaction generated by phorid defence behaviour. 3 In this study we test for the existence of these trade-offs and evaluate the predictions of a model for how they interact in an assemblage of woodland ants in which two behaviourally dominant members are attacked by phorid fly parasitoids as they attempt to harvest food resources. 4 The major findings are that unparasitized species in the assemblage follow a dominance–discovery trade-off curve. When not subject to attack by phorid flies, host species violate that trade-off by finding resources too quickly for their level of behavioural dominance. In contrast, when attacked by their phorid parasitoids, the host species dominance drops such that they fall into the assemblage trade-off. 5 These results match the predictions of the balance of terror model, which derives the optimal host response to parasitism, indicating that the host species balance the competing fitness costs of reduced competitive dominance and loss of workers to parasitism. This result supports the view that understanding the structure of ecological communities requires incorporating the indirect effects created by trait plasticity.

The ability to detoxify defensive compounds of competitors provides key ecological advantages that can influence community-level processes. Although common in plants and bacteria, this type of detoxification interaction is extremely rare in animals. Here, using laboratory behavioral assays and analyses of videotaped interactions in South America, we report widespread venom detoxification among ants in the subfamily Formicinae. Across both data sets, nine formicine species, representing all major clades, used a stereotyped grooming behavior to self-apply formic acid (acidopore grooming) in response to fire ant (Solenopsis invicta and S. saevissima) venom exposure. In laboratory assays, this behavior increased the survivorship of species following exposure to S. invicta venom. Species expressed the behavior when exposed to additional alkaloid venoms, including both compositionally similar piperidine venom of an additional fire ant species and the pyrrolidine/pyrroline alkaloid venom of a Monomorium species. In addition, species expressed the behavior following exposure to the uncharacterized venom of a Crematogaster species. However, species did not express acidopore grooming when confronted with protein-based ant venoms or when exposed to monoterpenoid-based venom. This pattern, combined with the specific chemistry of the reaction of formic acid with venom alkaloids, indicates that alkaloid venoms are targets of detoxification grooming. Solenopsis thief ants, and Monomorium species stand out as brood-predators of formicine ants that produce piperidine, pyrrolidine, and pyrroline venom, providing an important ecological context for the use of detoxification behavior. Detoxification behavior also represents a mechanism that can influence the order of assemblage dominance hierarchies surrounding food competition. Thus, this behavior likely influences ant-assemblages through a variety of ecological pathways.

Interaction modifications arise when the presence of one species alters the behavior of a second thereby altering that species' interactions with a third. Species-specific phorid parasitoids that attack ants at food resources can modify the competitive interactions between their host and competing ant species. This study examines whether interaction modifications created during interactions between the phorid fly parasitoid, Apocephalus sp. 8, its host, Pheidole diversipilosa, and ant species with which P. diversipilosa competes determine P. diversipilosa's foraging success. We also explore how varying resource size affects the ability of P. diversipilosa to capture resources and if this factor changes the nature or magnitude of the interaction modifications. We find that conflict between the host ant and competitor species at resources greatly improves the ability of the parasitoid to locate its host. In addition, the presence of a parasitoid at a food resource strongly reduces the ability of the host ant to capture that food resource when competing with other ant species. A wide variety of competitor species benefit from these linked interaction modifications and do so in accordance with their abundance in the assemblage. Finally, the size of the food resource in dispute affects the ability of the host ant to capture the resource, but parasitism and resource size affect resource turnover independently. These results are relevant to understanding the importance of interaction modifications in determining structure of ant assemblages.

A new microsporidian genus and species, Myrmecomorba nylanderiae, is described from North American populations of the tawny crazy ant, Nylanderia fulva. This new species was found to be heterosporous producing several types of binucleate spores in both larval and adult stages and an abortive octosporoblastic sporogony in adult ants. While microsporidia are widespread arthropod parasites, this description represents only the fifth species described from an ant host. Molecular analysis indicated that this new taxon is phylogenetically closely allied to the microsporidian family Caudosporidae, a group known to parasitize aquatic black fly larvae. We report the presence of 3 spore types (Type 1 DK, Type 2 DK, and octospores) with infections found in all stages of host development and reproductive castes. This report documents the first pathogen infecting N. fulva, an invasive ant of considerable economic and ecological consequence.

Know Your Enemy Fire ants accidentally introduced to North America from their native range in Argentina have been hugely invasive and difficult to eradicate and caused both environmental and economic damage. Recently, another accidentally introduced Argentine ant, the tawny crazy ant, appears to be displacing the fire ants. How? LeBrun et al. (p. 1014 , published online 13 February; see the Perspective by Kaspari and Weiser ) show that tawny crazy ants have a chemical and behavioral response to the toxic bite of fire ants that vastly reduces their mortality during confrontations and that allows the tawny crazy ants to outcompete their rivals.

Proctolaelaps kirmsei (Mesostigmata: Ascidae) is a specialist flower mite that feeds on pollen and nectar in flowers of Hamelia patens (Rubiaceae). We offered mites a choice between pairs of manipulated H. patens flowers in petri dishes in the laboratory, then counted the number of mites inside the flowers after 1 hour. Relative to flowers with stamens and nectaries removed, there were more mites in flowers with only stamens, with only nectaries, and with both stamens and nectaries. When offered a choice between flowers with stamens but no nectaries versus flowers with nectaries but no stamens, the mites did not show a preference. Mites preferred H. patens flowers with H. patens stamens over those with the stamens of a coexisting non-host plant (Heliconia pogonantha; Heliconiaceae) inserted, so the identity of the pollen is important in flower choice. These results and previous work on the importance of nectar show that the presence of both pollen and nectar can be used by P. kirmsei to discrimin...

Biological invasions represent communities in flux. Although stasis is never the rule in nature, biological interactions in communities usually occur within a framework of shared ecological and evolutionary history. Consequently, biological invasions represent unique opportunities to study dynamics that can otherwise be difficult to observe (Elton 1958). Invasive ants are excellent organisms with which to pursue this goal, in part because ants as a group play a variety of important ecological roles within biological communities (Holldobler and Wilson 1990). Ant invasions hold much potential for improving an understanding of ecological processes in general, as well as of interactions more specific to myrmecology. For instance, the dynamics that exist during ant invasions may reveal the traits that promote behavioural or ecological dominance. Furthermore, highly successful invasive ant species are often less conspicuous in their native ranges, so identification of the factors responsible for their greater prominence in introduced areas can provide insight into the more typical workings of ant communities, may highlight intrinsic differences between communities that have formed in separate biogeographic regions, and may suggest possible methods of control. Understanding ant invasion processes and causes of success thus has both basic and applied relevance. Human activities have introduced many ant species to new biogeographic regions (McGlynn 1999b; Chapter 13). While most introduced ant species have limited success in spreading away from the human-modified habitats in which they usually first arrive, a subset can invade nearby, or even distant, undisturbed natural habitats. This distinction is not always hard and fast, as virtually all species exhibit different degrees of invasiveness and ecological dominance in different locales, and sometimes even under similar conditions in the same locale (e.g. Abbott et al. 2007). Notwithstanding, the following species are the most consistent in their ability to penetrate natural ecosystems and affect the composition or abundance of native species within them (Holway et al. 2002a): the Argentine ant (Linepithema humile), the red imported fire ant (Solenopsis invicta), the tropical fire ant (S. geminata), the big-headed ant (Pheidole megacephala), the little fire ant (Wasmannia auropunctata), and the yellow crazy ant (Anoplolepis gracilipes) (see Plate 14). We focus on these species because of their pronounced invasive tendencies, and also because they have been studied the most intensively, especially with regard to the factors that control spread and underlie dominance. In fact, the majority of our knowledge on these topics comes from studies on just two species: S. invicta and L. humile. This represents both a major weakness and a clear avenue for progress in the field, pointing to a need for more complete information on the ecologies of other invasive ant species, as well as their close relatives that fail to become invasive (Chapter 13; see also Section 14.4). This limitation must be acknowledged at the outset, as it impinges on our under

Species abundances in natural systems are usually close to some equilibrium, making mechanisms that maintain or prevent species coexistence difficult to discern. Biological control projects provide an opportunity to observe systems transition between equilibriums as a result of the influence of the newly introduced species. In the southeastern United States and Texas, species of phorid fly parasitoids are being sequentially introduced as control agents for imported fire ants. The first two species introduced, Pseudacteon tricuspis and P. curvatus, partition the host niche based upon body size and co-exist broadly in their native range in Argentina, indicating they would form a co-existing and complementary suite of parasitoids in North America. This study examines the interaction between these parasitoids at multiple temporal and spatial scales. Surprisingly, data at all scales reveal that as P. curvatus establishes at a site it competitively displaces P. tricuspis. However, the speed of this reduction appears to differ between ecoregions, suggesting that the rate of displacement depends on environment. At the site where P. curvatus has been established the longest, this population interaction approaches complete displacement. Tests of potential mechanisms causing this displacement reveal that direct competition for host workers alters the operational sex ratio of the P. tricuspis population, but the strength of this effect is insufficient to explain the displacement. Experiments reveal the operation of a strong, indirect effect whereby locally common species preempt reproductive opportunities from rarer species by inducing host behavioral defenses. Finally, a re-examination of published data from their native range reveals that a previously overlooked negative relationship between the densities of these two species also exists there, suggesting that the same processes as those reported here also operate in South America.

Although the ecological effects of invasions often become obvious soon after introduced species become established, more gradual effects may take years to manifest and can thus require long-term data for quantification. We analyzed an 8-year record of stable isotope data on Argentine ants ( Linepithema humile ) from southern California to infer how the trophic position of this widespread invasive species changes over time as native ant species are displaced. We couple this longitudinal analysis with a biregional comparison of stable isotope data (δ 15 N) on ants from Argentina (native range) and California (introduced range) to quantify ( i ) how the trophic position of L. humile differs between native and introduced populations, and ( ii ) how relative trophic position as estimated by δ 15 N values of Argentine ants compare with those of other ants at the same site. Both long-term and biregional comparisons indicate that the Argentine ant's relative trophic position is reduced at sites with a longer history of occupation. Over the course of 8 years, the relative trophic position of L. humile remained high at the leading edge of an invasion front but declined, on average, behind the front as native ants disappeared. Relative to native populations, where L. humile is among the most carnivorous of ants, Argentine ants from California occupied lower trophic positions. These results support the hypothesis that Argentine ants shift their diet after establishment as a result of resource depletion and increasing reliance on plant-based resources, especially honeydew-producing Hemiptera. Our results demonstrate the value of long-term and biregional data in uncovering ecological effects of invasions.

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.

Ant communities often consist of many species with apparently similar niches. We present a mathematical model of the dominance-discovery trade-off, the trade-off between the abilities to find and to control resources, showing that it can in principle facilitate the coexistence of large numbers of species. Baiting studies of dominance and discovery abilities in an ant community from the Chiricahua Mountains of Arizona indicate that real communities fail to fit the assumptions of the simple model in several ways: (1) dominance depends on the size of the food resource; (2) for some ants, dominance depends on the presence or absence of specialist parasitoids; (3) pairwise dominance is not an all-or-nothing trait; and (4) a consistent negative relationship between pairwise differences in per capita discovery rates and dominance can be detected for only one bait type. Extended models incorporating these factors successfully predict the coexistence of five of the six most abundant members of this community but fail to accurately predict their relative abundances. Sensitivity analysis indicates that each complicating factor enhances the extent of coexistence.

A wide variety of animal communities are organized into interspecific dominance hierarchies associated with the control and harvest of food resources. Interspecific dominance relationships are commonly found to be linear. However, dominance relations within communities can form a continuum ranging from intransitive networks to transitive, linear dominance hierarchies. How interference competition affects community structure depends on the configuration of the dominance interactions among the species. This study explores how resource size and the trait-mediated indirect effect (TMIE) specialist phorid fly parasitoids exert on interference competition, affect the transitive nature of competitive interactions in an assemblage of woodland ants. I quantify the linearity of networks of interactions associated with large and small food resources in the presence and absence of phorid parasitoids. Two distinct, significantly linear dominance hierarchies exist within the ant assemblage depending on the size of the disputed resource. However, the presence of phorid fly parasitoids eliminates the linearity of both dominance hierarchies. The host's phorid defense behaviors reduce the competitive asymmetries between the host and its subdominant competitors, increasing the indeterminacy in the outcome of competitive interactions. Thus, both resource size variation and phorid-induced TMIEs appear to facilitate coexistence in assemblages of scavenging ants.

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.