Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

Combined effect of hemipteran control and liquid bait on Argentine ant populations

The invasive Argentine ant, Linepithema humile (Mayr), has become a worldwide problem capable of inflicting significant ecological and economic injury on urban, agricultural, and natural environments. The mobility of this pest ant has long been noted, rapidly moving nests to new food resources and then away as resources are depleted. This ant, like many pest ant species, has a special affinity for honeydew excreted by phloem-feeding Hemiptera. We investigated the effect of various hemipteran control strategies on terrapin scale densities and measured their indirect effect on local Argentine ant densities and foraging effort. We then determined whether this indirect treatment strategy improved the performance of an ant bait. We predicted that Argentine ants would move nests away from trees treated for Hemiptera and then move nests back when a liquid bait was offered, followed by a decline in ant numbers due to intake of the toxicant. A horticultural oil spray and soil application of the systemic insecticide, imidacloprid, had no effect on terrapin scale numbers. However, trunk-injected dicrotophos caused a reduction in scale and a decline in local Argentine ant nest density and canopy foraging effort. We also recorded a reduction in local Argentine ant ground foraging when large amounts of liquid bait were applied, and we found no evidence that combining dicrotophos with liquid ant bait performed better than each treatment alone. We suggest that a strategy of combined hemipteran control plus application of liquid ant bait can reduce local Argentine ant densities, when both components of this system are highly efficacious.     

Assessment of the Argentine ant invasion management by means of manual removal of winter nests in mixed cork oak and pine forests

The dynamic in Argentine ant colonies varies seasonally, influenced by biotic and abiotic factors. In winter the spatial range of the colony is contracted in large formations (winter nests) containing a large number of queens and workers. Winter nests are the clue to the species’ dispersion power and the invasion of new habitats. For this reason a yearly elimination of queens and workers in winter at the edge (front) of the invasion could be a useful tool for weakening the species’ dispersion and therefore limiting its establishment in new areas. Here, we determined the spatial dynamics of the Argentine ant nests during 1 year, and we assessed the invasion management by means of manual removal of winter nests for two consecutive winters, determining its effects during the following 3 years. We mapped nests found in 18 plots divided into two groups: extirpated (with removal of nests) and non-extirpated (control), along the fronts of three locations. Seasonal variation in the abundance of nests and workers, together with the two-year extirpation effects were evaluated. We found that colonies tended to follow an annual cycle of contraction and dispersion, with a decrease in the number of nests as we approached the invasion front. The extirpation was effective only in the front area, where it promoted smaller, less lasting and aggregated nests, as well as a decrease in the abundance of queens and workers. Nests also experienced a decrease during the two first winters but a recovery in the third, when no extirpation was done. Thus, a yearly perturbation should be performed to keep the expansion of the Argentine ant at a low rate, and to limit its establishment in new areas.     

Seasonal spatial dynamics and causes of nest movement in colonies of the invasive Argentine ant (Linepithema humile)

Abstract.  1. Colony organisation and movement behaviour of the Argentine ant ( Linepithema humile ) was studied over 3 years in field populations in California and in captive colonies in the laboratory. This invasive species is highly polydomous and unicolonial; colonies consist of expansive and fluid networks of nests and trails. The spatial and temporal organisation of colonies may contribute to ecological dominance.
2. Argentine ant nests and inter-nest trails shift in size, abundance, and location, so that colony networks are spatially contracted in the winter and expanded spring to autumn. Colonies occupy permanent sites; ants migrated to and from the same winter nest locations year after year, and occupied 30% of the same nests repeatedly during seasonal migrations.
3. Nests were moved on average 2–3 m. Forty-two per cent were occupied less than 1 month, 4% the entire study, and the other 54% lasted 3.9 ± 2.3 months (mean ± SD).
4. Nests were located within 2–4 m of woody plants, in warm sites in the winter and cool sites in the summer. Both humidity and food availability influenced nest-site choice in laboratory colonies. However, when faced with a trade-off between factors, the ants chose humid nest boxes over nest boxes near food, and ants moved nests only in response to changes in humidity and not distance to food.
5. The results indicate that L. humile colonies are seasonally polydomous, and that nest movements are driven by changes in microclimate. Colony organisation maintains high local density and increases food supply, which may improve the competitive ability of L. humile colonies and reduce opportunities for species coexistence.     

Social Control over the Survival and Selection of Winged Virgin Queens in an Ant without Nuptial Flight: Iridomyrmex humilis

Laboratory and field experiments show that the number of winged virgin queens of the Argentine ant Iridomyrmex humilis allowed to remain alive in the nest is under social control. When reared in their natal nest, survival of virgin queens depends on the presence of males. In the presence of mature alate males or even male pupae only 20% of the gynes are executed before they are 4 days old. By contrast, if pupal and adult males are absent, about 55% of the gynes are executed. The workers tend to kill the lightest ones. Although the Argentine ant is a unicolonial species exhibiting complete acceptance of foreign conspecific workers and mated queens, regardless of whether the recipient nest is queenright or queenless, the acceptance of gynes is variable: they are executed by foreign queenless workers but widely accepted in queenright nests. These results are discussed with regard to possible existence of queen and gyne pheromones allowing recognition and to the lack of nuptial flight in this species.     

Factors influencing the distribution of nests of the Argentine ant Linepithema humile Mayr (hymenoptera: formicidae), in a foothills ecosystem of the central zone of Chile

The purpose of this research was to demonstrate the dependant relationship between the Argentine ant Linepithema humile Mayr, plant coverage, and human settlements. A method was designed in order to sample the presence of the Argentine ant and other ant species that may had influenced L. humile distribution, under different types of vegetation and percentages of plant coverage, taking in consideration the distance from human settlements. Eight sample stations were established. Four of these stations were found on a dwelling home and the other four were found on similar areas, but far from human settlements. Three transects were established from the center of each sampling station (Tt1, shrublike vegetation, Tt2, herblike vegetation and Tt3, no vegetation). Transects were compound by 10 m side quadrants. A negative correlation was found between nest number of L. humile and human settlement distance, therefore nests were not found at distances higher than 80 m, in all studied areas. There was a positive and significant correlation between nest number of L. humile and plant coverage percentage, regardless the type of vegetation found. The same trend was found for other ant species studied in the area. Finally, it was proved that the distribution of L. humile nests is not influenced by other ant species.     

The Argentine ant persists through unfavorable winters via a mutualism facilitated by a native tree

Mutualisms and facilitations can fundamentally change the relationship between an organism's realized and fundamental niche. Invasive species may prove particularly suitable models for investigating this relationship as many are dependent on finding new partners for successful establishment. We conducted field-based experiments testing whether a native tree facilitates the successful survival of the invasive Argentine ant, Linepithema humile (Mayr), through unfavorable winter conditions in the southeastern United States. We found Argentine ant nests aggregated around the native loblolly pine, Pinus taeda L., during the winter months. The bark of this tree absorbed enough radiant solar energy to reach temperatures suitable for Argentine ant foraging even when ambient temperatures should have curtailed all foraging. Conversely, foraging ceased when the trunk was shaded. The sun-warmed bark of this tree gave the Argentine ant access to a stable honeydew resource. Argentine ants were not found on or near deciduous trees even though bark temperatures were warm enough to permit Argentine ant foraging on cold winter days. Augmenting deciduous trees with sucrose water through the winter months lead to Argentine ant nests remaining at their base and Argentine ants foraging on the tree. The Argentine ant requires both foraging opportunity and a reliable winter food source to survive through unfavorable winter conditions in the southeastern United States. The loblolly pine provided both of these requirements extending the realized niche of Argentine ants beyond its fundamental niche.     

Utilization of harvester ant nest sites by Persian goitered gazelle in steppes of central Iran

Ants are among the most important elements in many ecosystems and known as famous ecosystem engineers. By changing physical and chemical properties of soil, ants may provide suitable habitats for other species. Based on previous observations, we hypothesized that Persian goitered gazelles (Gazella subgutturosa subgutturosa) exhibit a preference for utilizing sites close to seed harvester ant (Messor spp.) nests. We tested our hypothesis by (1) mapping the occurrence of harvester ant nests and aggregated gazelle pellet groups along 31 strip transects, (2) monitoring pellet group accumulation bimonthly at 56 pairs of permanent plots established on ant nests and at adjacent control sites for a complete year, and (3) comparing vegetation and soil parameters between ant nest sites used by gazelles and paired control plots without ant nests. Although the area of Messor spp. nest sites covered only about 0.29% of the sampled transects, 84% of the gazelle pellet group aggregation sites were positioned upon ant nests, suggesting that gazelles actively selected Messor spp. nest sites. Pair-wise comparisons between ant nest plots and paired control plots also confirmed higher use of ant nest sites by gazelles compared to sites without ant nests in all time periods. Percent soil organic matter, percent cover of gravel, and annual herb vegetation significantly differed between ant nest and paired control plots in all the vegetation communities. We suggest that the alterations brought about by harvester ants on soil and vegetation make these sites attractive to gazelles. Gazelle territoriality behaviour and use of ant nests as bedding sites may be the reasons for selection of ant nest sites by gazelles.     

Plasticity in queen number and social structure in the invasive Argentine ant (Linepithema humile)

In many polygynous social insect societies, ecological factors such as habitat saturation promote high queen numbers by increasing the cost of solitary breeding. If polygyny is associated with constrained environments, queen number in colonies of invasive social insects should increase as saturation of their new habitat increases. Here I describe the variation in queen number, nestmate relatedness, and nest size along a gradient of time since colonization in an invading population of Argentine ants (Linepithema humile) in Haleakala, Hawaii. Nest densities in this population increase with distance from the leading edge of the invasion, reaching a stable density plateau approximately 80 m from the edge (> 2 years after colonization). Although the number of queens per nest in Haleakala is generally lower than previously reported for Argentine ants, there is significant variation in queen number across this population. Both the observed and effective queen numbers increase across the density gradient, and nests in the center of the population contain queen numbers three to nine times higher than those on the edge of the invasion. The number of workers per nest is correlated with queen number, and nests in the center are six times larger than nests at the edge. Microsatellite analysis of relatedness among nestmates reveals that all nests in the Haleakala population are characterized by low relatedness and have evidence of multiple reproducing queens. Relatedness values are significantly lower in nests in the center of the population, indicating that the number of reproducing queens is greater in areas of high nest density. The variation in queen number and nestmate relatedness in this study is consistent with expectations based on changes in ecological constraints during the invasion of a new habitat, suggesting that the social structure of Argentine ant populations is strongly influenced by ecological factors. Flexibility in social structure may facilitate persistence in variable environments and may also confer significant advantages to a species when introduced into new areas.     

Testing hypotheses about the function of repeated nest abandonment as a life history strategy in a passerine bird

Nest structures are essential for successful reproduction in most bird species. Nest construction costs time and energy, and most bird species typically build one nest per breeding attempt. Some species, however, build more than one nest, and the reason for this behaviour is often unclear. In the Grey Fantail Rhipidura albiscapa, nest abandonment before egg‐laying is very common. Fantails will build up to seven nests within a breeding season, and pairs abandon up to 71% of their nests before egg‐laying. We describe multiple nest‐building behaviour in the Grey Fantail and test four hypotheses explaining nest abandonment in this species: cryptic depredation, destruction of nests during storm events, and two anti‐predatory responses (construction of decoy nests to confuse predators, and increasing concealment to ‘hide’ nests more effectively). We found support for only one hypothesis – that abandonment is related to nest concealment. Abandoned nests were significantly less concealed than nests that received eggs. Most abandoned nests were not completely built and none received eggs, thus ruling out cryptic predation. Nests were not more likely to be abandoned following storm events. The decoy nest hypothesis was refuted as abandoned nests were constructed at any point during the breeding season and some nests were dismantled and the material used to build the subsequent nest. Thus, Grey Fantails are flexible about nest‐site locations during the nest‐building phase and readily abandon nest locations if they are found to have deficient security.     

Excavation and architecture of Argentine ant nests

We used x-ray computed tomography to study the elaboration of nest structures in small sand-filled nest boxes by Argentine ant (Linepithema humile) colonies composed of 10, 100, and 1000 workers. The pattern of nest growth was consistent with a process of density-dependent stimulation of excavation, which subsided as nests grew and the density of digging stimuli declined. Thus, nest excavation would be auto-regulating, and final nest size should be adjusted to colony size. We found that excavation rates and final nest sizes increased with colony size, but were not tenfold greater in 1000-worker colonies than in 100-worker colonies. In the largest colonies, the internal surface area scaled allometrically with volume, so that more surface was obtained relative to volume excavated as the nest grew. Although the gross features of Argentine ant nests, such as total size, seem explicable by a simple, self-organized regulatory process, other features of the nest architecture will require further investigation. Received 3 March 2005; revised 26 April 2005; accepted 3 May 2005.     

Beach erosion and nest site selection by the leatherback sea turtle Dermochelys coriacea (Testudines: Dermochelyidae) and implications for management practices at Playa Gandoca, Costa Rica

Leatherback sea turtles (Dermochelys coriacea) nest on dynamic, erosion-prone beaches. Erosive processes and resulting nest loss have long been presumed to be a hindrance to clutch survival. In order to better understand how leatherbacks cope with unstable nesting beaches, I investigated the role of beach erosion in leatherback nest site selection at Playa Gandoca, Costa Rica. I also examined the potential effect of nest relocation, a conservation strategy in place at Playa Gandoca to prevent nest loss to erosion, on the temperature of incubating clutches. I monitored changes in beach structure as a result of erosion at natural nest sites during the time the nest was laid, as well as in subsequent weeks. To investigate slope as a cue for nest site selection, I measured the slope of the beach where turtles ascended from the sea to nest, as well as the slopes at other random locations on the beach for comparison. I examined temperature differences between natural and relocated nest sites with thermocouples placed in the sand at depths typical of leatherback nests. Nests were distributed non-randomly in a clumped distribution along the length of the beach and laid at locations that were not undergoing erosion. The slope at nest sites was significantly different than at randomly chosen locations on the beach. The sand temperature at nest depths was significantly warmer at natural nest sites than at locations of relocated nests. The findings of this study suggest leatherbacks actively select nest sites that are not undergoing erosive processes, with slope potentially being used as a cue for site selection. The relocation of nests appears to be inadvertently cooling the nest environment. Due to the fact that leatherback clutches undergo temperature-dependent sex determination, the relocation of nests may be producing an unnatural male biasing of hatchlings. The results of this study suggest that the necessity of relocation practices, largely in place to protect nests from erosion, should be reevaluated to ensure the proper conservation of this critically endangered species.     

Rock selection for nesting inProformica longiseta Collingwood, 1978 (Hymenoptera,Formicidae) in a Mediterranean high mountain

Proformica longiseta, an ant species endemic to certain high mountains, has been studied with regard to its tendency to nest underneath rocks. This behaviour led us to investigate whether a particular type of rock is selected, and, if so, the advantages to this species of the selection. This study was located at 2400 m above sea-level in the Sierra Nevada mountains (Granada, Spain). To determine the range of rock sizes available, in eight control plots the thicknesses were recorded and also the maximum and minimum diameters of all rocks with surface area of more than 6 cm². A total of 1724 rocks were recorded, noting among the rock types available those that were used for nesting. Also as a preliminary estimate of the thermal characteristics of the rocks, we recorded the temperatures of the air, soil surface, soil under a rock covering an ant nest, and soil at 30 cm depth. The results indicate thatP. longiseta selects flat rocks with a surface area/thickness ratio of 2–5, the thickness being not less than 1.5 cm, the surface area being 50–250 cm² and volume less than 1000 cm³. This type of rock offers the nests two beneficial but apparently contradictory effects: (1) protection against high temperatures on hot days, and (2) warmth on colder days by absorbing the diffusing solar heat more effectively than the air or soil around the nest.     

The importance of pre-thicket conifer plantations for nesting Hen Harriers Circus cyaneus in Ireland

Hen Harriers Circus cyaneus are threatened across much of their range and their conservation requires appropriate habitat management. The locations of 148 Hen Harrier nests found in the Republic of Ireland during national breeding surveys in 2000 and 2005 were used to assess nest-site selection. The distribution of these nests was compared to distributions of randomly located points to investigate selection at the scale of the nest-site and landscape. The main nesting habitats selected were pre-thicket stage of first and, particularly, second rotation plantations, mostly of exotic conifers. There was no evidence that the area of post-closure plantations negatively affected Hen Harrier nest distribution. There was a positive correlation across study areas between changes in numbers of Hen Harrier nests between 2000 and 2005 and changes in the area of pre-thicket second rotation plantations over the same period. The overall effect of plantation forests on breeding Hen Harriers in Ireland therefore appears to be positive. However, this study did not consider the effects of plantation habitats on breeding success. Improved grassland was strongly avoided as a nesting habitat. Furthermore, after controlling for the influence of nesting habitat on nest location, landscapes with a high percentage cover of improved grassland were also avoided. Further agricultural intensification of grassland in areas where Hen Harriers breed is likely to have a negative impact on this species. These results are required for the development of management strategies for the conservation of this species.     

Ant Colonies Prefer Infected over Uninfected Nest Sites

During colony relocation, the selection of a new nest involves exploration and assessment of potential sites followed by colony movement on the basis of a collective decision making process. Hygiene and pathogen load of the potential nest sites are factors worker scouts might evaluate, given the high risk of epidemics in group-living animals. Choosing nest sites free of pathogens is hypothesized to be highly efficient in invasive ants as each of their introduced populations is often an open network of nests exchanging individuals (unicolonial) with frequent relocation into new nest sites and low genetic diversity, likely making these species particularly vulnerable to parasites and diseases. We investigated the nest site preference of the invasive pharaoh ant, Monomorium pharaonis, through binary choice tests between three nest types: nests containing dead nestmates overgrown with sporulating mycelium of the entomopathogenic fungus Metarhizium brunneum (infected nests), nests containing nestmates killed by freezing (uninfected nests), and empty nests. In contrast to the expectation pharaoh ant colonies preferentially (84%) moved into the infected nest when presented with the choice of an infected and an uninfected nest. The ants had an intermediate preference for empty nests. Pharaoh ants display an overall preference for infected nests during colony relocation. While we cannot rule out that the ants are actually manipulated by the pathogen, we propose that this preference might be an adaptive strategy by the host to “immunize” the colony against future exposure to the same pathogenic fungus.     

Ant behaviour and seed morphology: a missing link of myrmecochory

Seed dispersal by ants (myrmecochory) is mediated by the presence of a lipid-rich appendage (elaiosome) on the seed that induces a variety of ants to collect the diaspores. When seeds mature or fall onto the ground, these ant species transport them to their nest. After eating the elaiosome, the seed is discarded in nest galleries or outside, in the midden or farther away, where seeds can potentially germinate. The final location of seeds with their elaiosomes removed was evaluated to assess the importance of possible handles (structures that ants can grasp to carry) in transporting ants during re-dispersal experiments of seeds from nests of six species of ants. The results indicate that seeds remained within the nest because the ants were not able to transport them out of the nest. As a consequence of the elaiosome being removed, small ant species could not take Euphorbia characias seeds out of their nests. Only large ant species could remove E. characias seeds from their nests. Attaching an artificial handle to E. characias seeds allowed small ant species to redistribute the seeds from their nests. On the other hand, Rhamnus alaternus seeds that have a natural handle after the elaiosome removal were removed from the nests by both groups of ant species. If a seed has an element that acts as a handle, it will eventually get taken out of the nest. The ants’ size and their mandible gap can determine the outcome of the interaction (i.e. the pattern of the final seed shadow) and as a consequence, could influence the events that take place after the dispersal process.     

Nest-site selection and nest size influence the incubation temperature of Morelet's crocodiles

In oviparous tetrapods, the nesting-site selection by females is related to the habitat characteristics, which influences nest incubation temperature. Females can directly influence the incubation temperature by choosing certain construction materials or by building nests of different sizes. There are few studies focusing on these aspects in crocodilians that build mound nests. The aim of this study was to determine whether the nest size, its exposure to solar radiation, and the environmental temperature influence the incubation temperature of Morelet's crocodile (Crocodylus moreletii) nests. Artificial nests of two sizes (small and large) were constructed with similar characteristics to natural nests and placed in two locations differing in exposure to solar radiation (shaded and sunny). We used temperature and relative humidity data loggers to record the incubation temperature inside each nest every hour during the species' natural nesting period. Likewise, we recorded the ambient temperature every hour where the experiments were set up with temperature data loggers. We found that nest size and its exposure to solar radiation affected the incubation temperature, with smaller nests in shaded locations having lower incubation temperatures than larger nests in sunny locations. We discuss the importance of nest-site selection and maintenance behaviour of the mound nest by female crocodiles on the incubation temperature of the nest.     

Nest connectivity and colony structure in unicolonial Argentine ants

Unicolonial ant colonies occupy many nests and individuals rarely show aggression across large geographic distances. These traits make it difficult to detect colony structure. Here we identify colony structure at scales of hundreds of square-meters, within an invasive population of unicolonial Argentine ants. In experiments using labeled food, and in a 3-year census of nests and trails, we found that food was shared and nests were linked by trails at distances up to 50 meters. Food was not distributed to all nearby Argentine ant nests, showing that ants tend to share resources within a spatially bounded group of nests. The spatial extent of food sharing increased from winter to summer. Across different habitats and nest densities, nests were consistently aggregated at spatial scales of 3- 4 meters in radius. This suggests that new nests bud from old nests at short distances regardless of local conditions. We suggest that a ‘colony’ of Argentine ants could be defined as a group of nests among which ants travel and share food. In our study population, colonies occupy up to 650 m² and contain as many as 5 million ants. In combination with previous work showing that there is genetic differentiation among nests at similar spatial scales, the results suggest that Argentine ant populations do not function ecologically as single, large supercolonies, but instead as mosaics of smaller, distinct colonies consisting of groups of interacting nests. Received 6 June 2008; revised 30 June 2008; accepted 2 July 2008.     

Should I stay or should I go now? Patch use by African army ant colonies in relation to food availability and predation

Army ant colonies do not have permanent nests but frequently move to new patches. Local food depletion is considered the ultimate cause of this nomadic behaviour, but the proximate causes are not well understood. We tested if and how patch departure time of the aboveground-hunting army ant Dorylus molestus under field conditions is influenced by food availability and nest attacks by predators. In the first food supplement experiment, colonies receiving additional food throughout an entire nest stay did not reside in their nests for longer periods than control colonies. However, the distances travelled by colonies after nest stays during which colonies obtained food were shorter than those before these nest stays, indicating that colonies do assess food availability and avoid moving too far away from patches of high food availability. In the second food supplement experiment, in which colonies were given even larger amounts of food in the second half of their nest stay to mimic a rich unpredictable food source that these highly polyphagous predators are likely to encounter sometimes, patch departure times likewise did not differ between treated and control colonies. Either patch departure time is independent of food availability or there is another, as yet unappreciated proximate cause of colony movements in this species which we were unable to control for in our field experiments. One possibility is that encounters between neighbouring colonies influence patch departure time. In the experiment on the effect of predation, colonies responded to simulated nest attacks by mammals by leaving nests almost instantaneously and thus much earlier than control colonies. Rapid nest evacuation is likely a response to minimize the probability of repeat attacks by predators which cannot be repelled in other ways. Future studies will be necessary to definitively determine whether food availability influences patch departure times and to elucidate the consequences of colony encounters.     

Effects of experimental small-scale grassland fragmentation on spatial distribution, density, and persistence of ant nests

Abstract.  1. Grassland fragmentation is expected to influence the abundance of different invertebrate species to a different extent. Fragmentation-related effects are of particular importance in species that interact with many other species.
2. The density and spatial distribution of nests of 15 ant species in experimentally fragmented calcareous grasslands at three sites in the Northern Swiss Jura mountains were examined. Fragments of different size (0.25 m², 2.25 m², and 20.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland.
3. Three years after initiation of the experiment, ant nest density did not differ between fragments and control plots. Six years after initiation of the experiment, however, ant nest density and forager abundance were higher in large fragments than in large control plots. Ant nests tended to occur more frequently along the edge of fragments than in the core area. Persistence time of nests of the most abundant species, Lasius paralienus , tended to be shorter in fragments than in control plots. Furthermore, persistence time was longer in nests situated close to the fragment edge than in nests in the core area.
4. Effects on nest density, edge effects on the spatial distribution of nests, and the relationships between nest density and environmental factors were more pronounced when only nests of L. paralienus were considered. The implications of these findings for plant and other invertebrate species are discussed.