Behavioral interactions of the invasive Argentine ant with native ant species

Summary: The Argentine ant, Linepithema humile, has invaded many areas of the world, displacing native ants. Its behavior may contribute to its competitive success. Staged and natural encounters were observed at food resources in the field, between Argentine ants and eight ant species native to northern California. There was no relation between the frequency of aggression by any ant species and the outcome of encounters, though Argentine ants were more likely than ants of native species to behave aggressively. When an ant of one species initiated an encounter of any kind with an ant of another species, the ant that did not initiate was likely to retreat. This was true of all species studied. Most encounters between ants were initiated by Argentine ants. Thus the native species tended to retreat more frequently than Argentine ants. Interactions between Argentine ants and native species at food resources, causing ants of native species to retreat, may help Argentine ants to displace native species from invaded areas.     

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.     

Cryptic microsporidian parasites differentially affect invasive and native Artemia spp.

We investigated the host specificity of two cryptic microsporidian species (Anostracospora rigaudi and Enterocytospora artemiae) infecting invasive (Artemia franciscana) and native (Artemia parthenogenetica) hosts in sympatry. Anostracospora rigaudi was on average four times more prevalent in the native host, whereas E. artemiae was three times more prevalent in the invasive host. Infection with An. rigaudi strongly reduced female reproduction in both host species, whereas infection with E. artemiae had weaker effects on female reproduction. We contrasted microsporidian prevalence in native A. franciscana populations (New World) and in both invaded and non-invaded Artemia populations (Old World). At a community level, microsporidian prevalence was twice as high in native compared with invasive hosts, due to the contrasting host-specificity of An. rigaudi and E. artemiae. At a higher biogeographical level, microsporidian prevalence in A. franciscana did not differ between the invaded populations and the native populations used for the introduction. Although E. artemiae was the only species found both in New and Old World populations, no evidence of its co-introduction with the invasive host was found in our experimental and phylogeographic tests. These results suggest that the success of A. franciscana invasion is probably due to a lower susceptibility to virulent microsporidian parasites rather than to decreased microsporidian prevalence compared with A. parthenogenetica or to lower microsporidian virulence in introduced areas.     

Unadapted behaviour of native,dominant ant species during the colonization of an aggressive,invasive ant

Among the factors driving the invasive success of non-indigenous species, the “escape opportunity” or “enemy release” hypothesis argues that an invader’s success may result partly from less resistance from the new competitors found in its introduced range. In this study, we examined competitive interactions between the little fire ant Wasmannia auropunctata (Roger) and ant species of the genus Pheidole in places where both are native (French Guiana) and in places where only species of Pheidole are native (New Caledonia). The experimental introduction of W. auropunctata at food resources monopolized by the Pheidole species induced the recruitment of major workers only for the Guianian Pheidole species, which were very effective at killing Wasmannia competitors. In contrast, an overall decrease in the number of Pheidole workers and no recruitment of major workers were observed for the New Caledonian species, although the latter were the only ones able to kill the Wasmannia workers. These results emphasize the inappropriate response of native dominant New Caledonian species to W. auropunctata and, thus, the importance of enemy recognition and specification in the organization of ant communities. This factor could explain how invasive animal species, particularly ants, may be able to successfully invade species-rich communities.     

Modelling the potential geographic distribution of invasive ant species in New Zealand

Despite their economic and environmental impacts, there have been relatively few attempts to model the distribution of invasive ant species. In this study, the potential distribution of six invasive ant species in New Zealand are modelled using three fundamentally different methods (BIOCLIM, DOMAIN, MAXENT). Species records were obtained from museum collections in New Zealand. There was a significant relationship between the length of time an exotic species had been present in New Zealand and its geographic range. This is the first time such a time lag has been described for exotic ant species, and shows there is a considerable time lag in their spread. For example, it has taken many species several decades (40–60 years) to obtain a distribution of 17–25% of New Zealand regions. For all six species, BIOCLIM performed poorly compared to the other two modelling methods. BIOCLIM had lower AUC scores and higher omission error, suggesting BIOCLIM models under-predicted the potential distribution of each species. Omission error was significantly higher between models fitted with all 19 climate variables compared to those models with fewer climate variables for BIOCLIM, but not DOMAIN or MAXENT. Widespread species had a greater commission error. A number of regions in New Zealand are predicted to be climatically suitable for the six species modelled, particularly coastal and lowland areas of both the North and South Islands.     

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.     

Plant-based food resources,trophic interactions among alien species,and the abundance of an invasive ant

Recent research on invasive ants suggests that their success may be facilitated by increased resources at introduced locations stemming from the emergence of novel trophic interactions with abundant honeydew-producing Hemiptera. Moreover, those Hemiptera may themselves often be introduced or invasive. To test the importance of mutualisms for invasive species, we conducted a study in the southeastern United States of factors hypothesized to affect the abundance of an invasive ant native to South America, Solenopsis invicta. The study was conducted within grazing pastures, where S. invicta can be extremely abundant while also exhibiting substantial variability in abundance. A path analysis showed that the abundance of S. invicta was strongly and positively affected by the abundance of an invasive honeydew-producing mealybug native to Asia, Antonina graminis, and by the mealybugs’ host grasses because of their strong positive effect on mealybug abundance. Abundance of the mealybug was primarily attributable to an invasive host grass native to Africa, Cynodon dactylon. The abundance of S. invicta was also positively affected by the abundance of other arthropods that they are likely to consume, and those arthropods were positively affected by the abundance of both the A. graminis host grasses and other plants. Thus the study shows that the distribution and abundance of different plant species could have important effects on the abundance of S. invicta through their effect on the ants’ food resources. The results are also consistent with the hypothesis that the emergence of novel trophic interactions among invasive species can promote the abundance of invasive ants.     

Exploitative strategies of the invasive Argentine ant (Linepithema humile) and native ant species in a southern Spanish pine forest

The Argentine ant, Linepithema humile (Mayr, 1868), is displacing native ant species in Do?ana National Park (Spain). This paper discusses the results of experiments aimed at analyzing exploitation competition between the invading species and other ant species in a park community. The Argentine ant was found to implement several strategies favoring its success in exploitation competition: mass recruitment, use of various microhabitats (on the ground and in trees), and activity over a wide range of temperatures. Although these strategies were not exclusive to L. humile, their joint use, together with the large number of workers forming each "unicolony," conferred a clear advantage for resource exploitation. Some native species were more severely affected than others by the presence of L. humile in terms of both abundance and behavior. The worst affected species were those whose ecological characteristics were similar to those of the Argentine ant, e.g., Pheidole pallidula (Nylander, 1849); the species least affected was Cataglyphis floricola Tinaut, 1993, possibly because of its subordinate and thermophilous nature (little overlap of daily activity rhythms with the exotic species).     

Historic land use influences contemporary establishment of invasive plant species

The legacy of agricultural land use can have widespread and persistent effects on contemporary landscapes. Although agriculture can lead to persistent changes in soil characteristics and plant communities, it remains unclear whether historic agricultural land use can alter the likelihood of contemporary biological invasions. To understand how agricultural land-use history might interact with well-known drivers of invasion, we conducted factorial manipulations of soil disturbance and resource additions within non-agricultural remnant sites and post-agricultural sites invaded by two non-native Lespedeza species. Our results reveal that variation in invader success can depend on the interplay of historic land use and contemporary processes: for both Lespedeza species, establishment was greater in remnant sites, but soil disturbance enhanced establishment irrespective of land-use history, demonstrating that contemporary processes can help to overcome legacy constraints on invader success. In contrast, additions of resources known to facilitate seedling recruitment (N and water) reduced invader establishment in post-agricultural but not in remnant sites, providing evidence that interactions between historic and contemporary processes can also limit invader success. Our findings thus illustrate that a consideration of historic land use may help to clarify the often contingent responses of invasive plants to known determinants of invasibility. Moreover, in finding significantly greater soil compaction at post-agricultural sites, our study provides a putative mechanism for historic land-use effects on contemporary invasive plant establishment. Our work suggests that an understanding of invasion dynamics requires knowledge of anthropogenic events that often occur decades before the introduction of invasive propagules.     

Contrasting effects of an invasive ant on a native and an invasive plant

When invasive species establish in new environments, they may disrupt existing or create new interactions with resident species. Understanding of the functioning of invaded ecosystems will benefit from careful investigation of resulting species-level interactions. We manipulated ant visitation to compare how invasive ant mutualisms affect two common plants, one native and one invasive, on a sub-tropical Indian Ocean island. Technomyrmex albipes, an introduced species, was the most common and abundant ant visitor to the plants. T. albipes were attracted to extrafloral nectaries on the invasive tree (Leucaena leucocephala) and deterred the plant’s primary herbivore, the Leucaena psyllid (Heteropsylla cubana). Ant exclusion from L. leucocephala resulted in decreased plant growth and seed production by 22% and 35%, respectively. In contrast, on the native shrub (Scaevola taccada), T. albipes frequently tended sap-sucking hemipterans, and ant exclusion resulted in 30% and 23% increases in growth and fruit production, respectively. Stable isotope analysis confirmed the more predacious and herbivorous diets of T. albipes on the invasive and native plants, respectively. Thus the ants’ interactions protect the invasive plant from its main herbivore while also exacerbating the effects of herbivores on the native plant. Ultimately, the negative effects on the native plant and positive effects on the invasive plant may work in concert to facilitate invasion by the invasive plant. Our findings underscore the importance of investigating facilitative interactions in a community context and the multiple and diverse interactions shaping novel ecosystems.     

Changes in the photosynthetic characteristics of cotton leaves infested by invasive mealybugs tended by native ant species

Herbivore injury has a direct effect on the growth and performance of host plants through photosynthetic suppression. However, changes in plant photosynthesis affected by ant tending of hemipteran sap feeders remain poorly understood. We investigated the effects of an invasive mealybug (Phenacoccus solenopsis) tended by native ants (Paratrechina longicornis) on the chlorophyll content and chlorophyll fluorescence characteristics of cotton (Gossypium hirsutum) leaves under greenhouse conditions. The results showed that the relative chlorophyll content of the infested cotton leaves significantly decreased after 10 days, and the chlorophyll contents were reduced by 26.4 and 34.9 % after 20 days in the without and with ant treatments compared to the control, respectively. In addition, the light utilization efficiency and maximum relative electron transport rate were reduced by 53.0 and 51.3 % compared to the control, respectively. However, no significant differences in these factors were found between the without and with ant treatments. The light saturation coefficient, describing the capacity of a sample to resist glare, exhibited no significant difference among treatments. The number of tending P. longicornis ants increased with P. solenopsis numbers, and the P. solenopsis numbers decreased after 20 days compared to the without ant treatment. We suggest that the tending ants may enhance the feeding ability of individual mealybugs in spite of the decreased number of mealybugs in this situation. Additionally, P. longicornis decrease the relative chlorophyll content of infested cotton leaves and may accelerate the damage caused by P. solenopsis to plants over time.     

Similar biotic factors affect early establishment and abundance of an invasive plant species across spatial scales

Research in community invasibiliy has focused on biotic and abiotic factors that influence the establishment of invasive species and whether such factors vary with spatial scale. Here, we investigate the role of both biotic and abiotic factors associated with the initial establishment of Lespedeza cuneata (L. cuneata) and its abundance at three spatial scales: neighborhoods (9-m² plots), communities (50-m² transect) and old fields (5,000–70,000 m²). We asked: (1) Do resource availability and community structure affect the establishment of L. cuneata?, and (2) Are resource availability and community structure associated with patterns of L. cuneata abundance from neighborhood scales to old-field scales? To investigate the first question, we manipulated soil nitrogen (N) availability at three levels in an existing old-field community and tracked emergence and persistence of L. cuneata seedlings, as well as total plant biomass of the community, availability of light, and soil moisture content. To address the second question, we performed surveys in which we estimated L. cuneata foliar cover at community scales (50-m² belt transects) and old-field scales (total area of 28 ha), and assessed the same biotic and abiotic variables as in the field experiment. The experiment revealed that establishment and persistence by L. cuneata seedlings were 15× and 5× lower in N-added plots than in N-reduced plots. Total plant community biomass was 30% greater in N-added plots than in N-reduced plots. Conversely, light and soil moisture were 60 and 20% lower in N-added plots than in N-reduced plots. Surveys of old fields indicated that community biomass was positively associated with L. cuneata cover at old-field scales likely resulting from greater soil N input from nitrogen fixation in fields with greater L. cuneata cover. In sum, these results indicate that biotic factors associated with establishment of a Rank 1 invasive plant species at the community scale are also related to its distribution at the old-field scale, but the direction of such associations changed across scales.     

Surrounding land use significantly influences adult mosquito abundance and species richness in urban mangroves

Mangroves harbor mosquitoes capable of transmitting human pathogens; consequently, urban mangrove management must strike a balance between conservation and minimizing public health risks. Land use may play a key role in shaping the mosquito community within urban mangroves through either species spillover or altering the abundance of mosquitoes associated with the mangrove. In this study, we explore the impact of land use within 500 m of urban mangroves on the abundance and diversity of adult mosquito populations. Carbon dioxide baited traps were used to sample host-seeking female mosquitoes around nine mangrove forest sites along the Parramatta River, Sydney, Australia. Specimens were identified to species and for each site, mosquito species abundance, species richness and diversity were calculated and were analyzed in linear mixed effects models. We found that the percentage of residential land and bushland in the surrounding area had a negative effect on mosquito abundance and species richness. Conversely, the amount of mangrove had a significant positive effect on mosquito abundance, and the amount of industrial land had a significant positive effect on species richness. These results demonstrate the need for site-specific investigations of mosquito communities associated with specific habitat types and the importance of considering surrounding land use in moderating local mosquito communities. A greater understanding of local land use and its influence on mosquito habitats could add substantially to the predictive power of disease risk models and assist local authorities develop policies for urban development and wetland rehabilitation.     

Exotic thistles increase native ant abundance through the maintenance of enhanced aphid populations

Exotic species change the structure and composition of invaded communities in multiple ways, but the sign of their impact on native species is still controversial. We evaluated the effects of the thistles Carduus thoermeri and Onopordum acanthium—two of the most abundant exotic plant species in disturbed areas of the Patagonian steppe—on the native tending ant assemblage. Exotic thistles showed an increased number of plants with aphids and had greater aphid density than native plants. Since native tending ants were present only in plants with aphids, their abundance was higher in infested thistles than in native plants. Path analyses confirmed that ant activity depended more on aphid density than on thistle traits. Our results suggest that the presence of exotic thistles in disturbed areas of NW Patagonia indirectly benefit the native ant assemblage through the maintenance of an increased aphid population. This illustrates how the impact of exotic on native species can depend on the ecological context.     

Biogeographic comparisons of the traits and abundance of an invasive crab throughout its native and invasive ranges

High abundances of non-native species in the invaded range may be linked to changes in fitness related traits. However, few studies have compared differences in both life-history traits and abundances of introduced species between their native and invaded ranges. We determined differences in 12 morphological traits, an important fitness related trait (body size), and the abundance of the porcelain crab, Petrolisthes elongatus, in its native (New Zealand) and invasive (Tasmania, Australia) ranges. P. elongatus was more abundant in the introduced range; however, changes in abundance depended on tidal height, with higher abundances only at mid and low tidal zones. The biomass of male crabs was higher in the invaded range compared to the native range, but there was no difference in female biomass between ranges. Despite increases in male biomass, sex ratios between native and invasive populations did not differ. In addition, principal components analysis showed no differences in overall morphology between Tasmania and New Zealand. Our study indicates that increased abundance in the invaded range of P. elongatus may be linked to high values of an important trait (greater biomass) in the invaded range. Importantly, changes in biomass and abundance may be due to P. elongatus being able to utilise mid/low zones more in the invaded range. Furthermore, our findings indicate that understanding how sex specific changes in biomass interact with the recipient environment (biotic and abiotic) in the introduced range will be important for determining the mechanisms underpinning the establishment and spread of P. elongatus.     

The distribution of an invasive fish species is highly affected by the presence of native fish species: evidence based on species distribution modelling

Topmouth gudgeon (Pseudorasbora parva) is one of the most invasive aquatic fish species in Europe and causes adverse effects to ecosystem structure and functioning. Knowledge and understanding of the species’ interactions with the environment and with native fish are important to stop and prevent the further spread of the species. Creating species distribution models is a useful technique to determine which factors influence the occurrence and abundance of a species. We applied three different modelling techniques: general additive models, random forests and fuzzy habitat suitability modelling (FHSM) to assess the habitat suitability of topmouth gudgeon. The former two techniques indicated that the abundance of native fish (i.e. biotic variables) was more important than environmental variables when determining the abundance of topmouth gudgeon in Flanders (Belgium). Bitterling (Rhodeus amarus), stone loach (Barbatula barbatula), three-spined stickleback (Gasterosteus aculeatus) and predator abundance were selected as the most important biotic variables and implemented in the FHSM to investigate species interactions. Depending on the preferred food source and spawning behaviour, either coexistence or interspecific competition can occur with bitterling, stone loach and three-spined stickleback. In contrast, the presence of predators clearly had a top down effect on topmouth gudgeon abundance. These findings could be applied as a biological control measure and implemented in conservation strategies in order to reduce the abundance of earlier established populations of topmouth gudgeon.     

Habitat alteration increases invasive fire ant abundance to the detriment of amphibians and reptiles

Altered habitats have been suggested to facilitate red imported fire ant (Solenopsis invicta) colonization and dispersal, possibly compounding effects of habitat alteration on native wildlife. In this study, we compared colonization intensity of wood cover boards by S. invicta among four forest management treatments in South Carolina, USA: an unharvested control (>30 years old); a partially thinned stand; a clearcut with coarse woody debris retained; and a clearcut with coarse woody debris removed. Additionally, we compared dehydration rates and survival of recently metamorphosed salamanders (marbled salamanders, Ambystoma opacum, and mole salamanders, A. talpoideum) among treatments. We found that the number of wood cover boards colonized by S. invicta differed significantly among treatments, being lowest in the unharvested forest treatments and increasing with the degree of habitat alteration. Salamanders that were maintained in experimental field enclosures to study water loss were unexpectedly subjected to high levels of S. invicta predation that differed among forest treatments. All known predation by S. invicta was restricted to salamanders in clearcuts. The amount of vegetative ground cover was inversely related to the likelihood of S. invicta predation of salamanders. Our results show that S. invicta abundance increases with habitat disturbance and that this increased abundance has negative consequences for amphibians that remain in altered habitats. Our findings also suggest that the presence of invasive S. invicta may compromise the utility of cover boards and other techniques commonly used in herpetological studies in the Southeast.     

Water guns affect abundance and behavior of bigheaded carp and native fish differently

Water guns have shown the potential to repel nuisance aquatic organisms. This study examines the effects of exposure to a 1966.4 cm³ seismic water gun array (two guns) on the abundance and behavior of Bighead Carp Hypophthalmichthys nobilis, Silver Carp H. molitrix (collectively referred to as bigheaded carp) and native fishes (e.g., Smallmouth Buffalo Ictiobus bubalus). Water guns were deployed in a channel that connects the Illinois River to backwater quarry pits that contained a large transient population of bigheaded carp. To evaluate the effect of water guns, mobile side-looking split-beam hydroacoustic surveys were conducted before, during and between replicated water gun firing periods. Water guns did not affect abundance of bigheaded carp, but abundance of native fish detected during the firing treatment was 43 and 34% lower than the control and water guns off treatments, respectively. The proximity of bigheaded carp to the water gun array was similar between the water guns on and water guns off treatments. In contrast, the closest detected native fish were detected farther from the water guns during the water guns on treatment (mean ± SE, 32.38 ± 3.32 m) than during the water guns off treatment (15.04 ± 1.59 m). The water gun array had a greater impact on native fish species than on bigheaded carp. Caution should be taken to the extrapolation of these results to other fish species and to fish exposed to water guns in different environments (e.g., reduced shoreline interaction) or exposure to a larger array of water guns, or for use of water guns for purposes other than a barrier.