Leaf-cutting ant nests near roads increase fitness of exotic plant species in natural protected areas

Understanding the mechanisms that promote the invasion of natural protected areas by exotic plants is a central concern for ecology. We demonstrated that nests of the leaf-cutting ant, Acromyrmex lobicornis, near roadsides promote the abundance, growth and reproduction of two exotic plant species, Carduus nutans and Onopordum acanthium, in a national park in northern Patagonia, Argentina and determine the mechanisms that produce these effects. Refuse dumps (RDs) from ant nests have a higher nutrient content than nearby non-nest soils (NNSs); foliar nutrient content and their 15N isotopic signature strongly suggest that plants reach and use these nutrients. Both species of exotic plants in RDs were 50-600% more abundant; seedlings had 100-1000% more foliar area and root and leaf biomass; and adult plants produced 100-300% more seeds than nearby NNS plants. Plants can thus gain access to and benefit from the nutrient content of ant RD, supporting the hypotheses that enhanced resource availability promotes exotic plant performance that could increase the likelihood of biological invasions. The two exotics produce an estimated of 8385000 more seeds ha(-1) in areas with ant nests compared with areas without; this exceptional increase in seed production represents a potential threat to nearby non-invaded communities. We propose several management strategies to mitigate this threat. Removal efforts of exotics should be focused on ant RDs, where plants are denser and represent a higher source of propagules.     

Bottom-up effects of host-plant species diversity and top-down effects of ants interactively increase plant performance

While plant diversity is well known to increase primary productivity, whether these bottom-up effects are enhanced by reciprocal top-down effects from the third trophic level is unknown. We studied whether pine tree species diversity, aphid-tending ants and their interaction determined plant performance and arthropod community structure. Plant diversity had a positive effect on aphids, but only in the presence of mutualistic ants, leading to a threefold greater number of both groups in the tri-specific cultures than in monocultures. Plant diversity increased ant abundance not only by increasing aphid number, but also by increasing ant recruitment per aphid. The positive effect of diversity on ants in turn cascaded down to increase plant performance; diversity increased plant growth (but not biomass), and this effect was stronger in the presence of ants. Consequently, bottom-up effects of diversity within the same genus and guild of plants, and top-down effects from the third trophic level (predatory ants), interactively increased plant performance.     

Native harvester ants threatened with widespread displacement exert localized effects on serpentine grassland plant community composition

Seed-harvesting ants can influence the abundance and distribution of plant species through both the selective harvesting of seeds and the construction of nutrient-rich nest mounds, but the relative contributions of these two mechanisms have not been addressed by previous studies. Furthermore, the impact of ant seed harvesting in California serpentine grasslands remains unresolved because of divergent results from several previous experiments. This study investigates the influence of harvester ants on serpentine grassland plant species composition by examining two potential signatures of seed harvesting ants on plant community composition: species composition on versus off ant nest mounds, and species abundance as a function of distance from nest mounds. Of the 28 plant species identified in this study, 22 exhibited spatial patterns consistent with effects of seed harvesting, nest construction, or both. Although most species showed significant gradients in abundance with distance from a nest, there were no clear relationships between plant species distributions and previously reported harvester ant seed foraging patterns. Harvester ant nest mounds supported plant communities that were distinct from the surrounding serpentine grassland, with notably higher densities of legumes and invasive annual grasses. Comparison of our results with those of previous studies indicates that the patterns we observed are generally consistent over time, but affect more species and a larger fraction of the grassland than previously reported. Unaffected areas of the grassland seem likely to serve as important refuges for some plant species.     

Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation.We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production.We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.     

Effects of harvester ants on plant species distribution and abundance in a serpentine grassland

Seed harvesting ants can have important effects on the composition and structure of plant communities. We investigated two effects of Messor andrei, the black seed-harvesting ant, on a serpentine grassland plant community in northern California. First, to determine if selective seed predation by ants affects plant community composition, we excluded harvester ants from 1-mediameter circular plots of grassland. Abundances of all species on these plots and on control plots were measured before and after exclosure. Second, to determine if M. andrei nest mounds affect plant community composition, we compared plant species abundances on and off nest mounds. M. andrei deposit large amounts of organic matter on their nest mounds over a foraging season, so mounds may alter the edaphic environment. The exclusion of seed-harvesting activity did not cause changes in the plant community. Nest mounds had a strong effect on plant communities: there were many more grasses and fewer forbs on ant mounds, although at least one forb, Lepidium nitidum, produced twice as many seeds when it grew on nest mounds. We found that nest mounds formed islands of higher-temperature soil in the serpentine grassland. Received: 31 March 1997 / Accepted: 6 May 1997     

Effects of Formica podzolica ant colonies on soil moisture,nitrogen, and plant communities near nests

1. Ants are widely regarded as ‘ecosystem engineers’ because their nest construction and contributions to nutrient cycling change the biological, chemical, and physical properties of the soil around their nests. Despite increasing attention to ant manipulation of soil ecosystems, the extent to which many common species influence soil properties, as well as nutrient uptake and community composition of plants near nests, is still unknown. 2. This study tested hypotheses that activities of a common subalpine ant, Formica podzolica, alter soil moisture and pH, redistribute nitrogen around nests, and affect plant species abundance and ground cover. 3. A combination of field sampling techniques showed that distance from a nest had a positive relationship with soil moisture and a negative relationship with plant abundance next to and downhill from nests. Slope aspect also affected plant communities, with downhill transects having higher plant cover and above‐ground biomass than uphill transects. A stable isotope analysis did not reveal that plants near nests had enriched ¹⁵N, but there were substantial differences in ¹⁵N among sites. 4. Overall, this study uncovers significant impacts of F. podzolica on the subalpine microhabitats directly surrounding their nests.     

Foliage-dwelling ants in a neotropical savanna: effects of plant and insect exudates on ant communities

Ant dominance in tropical ecosystems can be explained by a capacity to exploit liquid foods such as extrafloral nectaries (EFNs) and secretions from honeydew-producing hemipterans (HPHs). Such nutritious exudates may determine ant distribution in space and shape specialization in ant–plant interactions. We provide a first assessment of how EFNs and HPHs mediate the structure of ant assemblages, ant visitation intensity, and characteristics of ant–plant interaction networks across space in Brazilian “cerrado” savanna. We used arboreal pitfall traps to sample visiting ants in four cerrado localities and recorded the presence of lepidopteran larvae to determine their possible response to ant visitation. Ant species composition and richness did not differ regardless of the presence of liquid rewards on plants, and most network patterns did not show consistent differences. However, in two of the four sites, ant densities were higher on plants with HPHs or EFNs due to increased activity by Camponotus and Pseudomyrmex ants. At these two sites, plants with liquid food sources had a more specific ant assemblage (higher specialization d′) than did plants without resources, and caterpillars were more frequently found on plants with fewer workers of Camponotus and Pseudomyrmex. Plants with HPHs had increased ant visitation and accumulated more ant species than did plants with EFNs or without liquid foods. Ant response to such food sources may thus depend on local conditions and identity of ant species, and may determine how ant assemblages are structured. Results highlight how different patterns of ant visitation to liquid resources can produce distinctive effects on herbivore infestation.     

Genetic structure and reproductive strategy of the ant Cardiocondyla elegans: strictly monogynous nests invaded by unrelated sexuals

Cardiocondyla elegans is a Mediterranean ant that nests on river banks. It rears only wingless (ergatoid) males that live peacefully in the same nest as opposed to other species of the same genus, which have both peaceful, winged and mutually aggressive 'ergatoid' males. Using microsatellite analysis, we investigated the genetic structure of 21 colonies from three different locations as well as the parentage of sexuals of two colonies of C. elegans. We show that C. elegans is strictly monogynous, and that its nests can contain foreign sexuals. The presence of alien sexuals inside ant nests is described for the first time and probably counteracts inbreeding resulting from matings between siblings. In the laboratory, aggression tests showed that workers only allow alien males to enter their nests, while all winged female sexuals attempting to enter were attacked. Nevertheless, the presence of alien female sexuals in nests in the field seems to result from active carrying behaviour by workers during the reproductive period.     

几种杀虫剂对热带雨林生态区红火蚁蚁巢和工蚁的灭杀效果

为筛选出适用于云南热带雨林生态区的红火蚁Solenopsis invicta Buren防治药剂,通过7种杀虫剂对红火蚁蚁巢田间灭除效果试验测定。结果表明,20%多杀霉素悬浮剂（SC）、30%螺虫·噻虫嗪悬浮剂（SC）、70%吡虫啉可湿性粉剂（WP）、10%溴虫氟苯双酰胺悬浮剂（SC）和1%联苯·噻虫嗪颗粒剂（GR）对红火蚁蚁巢处理效果较好,药后1 d和10 d虫口减退率分别在95%和90%以上。其中10%溴虫氟苯双酰胺SC和1%联苯·噻虫嗪GR可作为红火蚁巢点处置的首选药剂;0.05%茚虫威饵剂（EB）速效性差,但药后10 d虫口减退率上升到86.8%。1.8%阿维菌素乳油（EC）药后1 d和10 d虫口减退率分别为89.3%和81%。这2种药剂可与前5种药剂交替使用。     

A field assessment of optimal foraging in ants: trail patterns and seed retrieval by the European harvester ant Messor barbarus

Summary: The ant Messor barbarus is a major seed predator on annual grasslands of the Mediterranean area. This paper is an attempt to relate the foraging ecology of this species to resource availability and to address several predictions of optimal foraging theory under natural conditions of seed harvesting.¶Spatial patterns of foraging trails tended to maximise acquisition of food resources, as trails led the ants to areas where seeds were more abundant locally. Moreover, harvesting activity concentrated on highly frequented trails, on which seeds were brought into the nest in larger numbers and more efficiently, at a higher mean rate per worker.¶The predictions of optimal foraging theory that ants should be more selective in both more resource-rich and more distant patches were tested in the native seed background. We confirm that selectivity of ants is positively related to trail length and thus to distance from the nest of foraged seeds. Conversely, we fail to find a consistent relationship between selectivity and density or species diversity of seed patches. We discuss how selectivity assessed at the colony level may depend on factors other than hitherto reported behavioural changes in seed choice by individual foragers.     

Influence of nests of leaf‐cutting ants on plant species diversity in road verges of northern Patagonia

Abstract. It has been suggested that ant nests are the most frequent small‐scale disturbance that affect vegetation patterns. However, their effects on plant diversity are little studied. We document effects of nests of the leaf‐cutting ant Acromyrmex lobicornis on physical‐chemical soil properties and their influence on plant diversity near road verges in a desert steppe in NW Patagonia, Argentina. We analysed nest soils and controls for nitrogen, phosphorus, organic matter, moisture retention capacity and texture. We also analysed the vegetation on 42 nests (30 active and 12 abandoned or without life) and 42 areas without nests. Soil around nests had a greater nutrient content and capacity to retain moisture than control soils, which is mainly due to the presence of organic waste that the ants deposit on the soil surface. We found no association between the occurrence of nests and specific groups of plants, but plant diversity was higher at nest‐sites than at nearby non‐nest sites. This increased diversity – which is also found on abandoned nests – is mainly due to the occurrence of a larger number of native and exotic plant species on nest‐sites that are uncommon elsewhere in the study area. The most abundant plant species showed similar cover values at nest and non‐nest sites. This suggests that changes in diversity are associated to edaphic changes caused by nests rather than by changes in competitive balance caused by dominant species exclusion. We propose that the nests of Acromyrmex lobicornis, through increasing the availability of resources, generate favourable microsites that can function both as ‘refuges’ for less frequent native species, and as‘stepping stones’ for less frequent exotic plant species.     

Ant-seed interactions: combined effects of ant and plant species on seed removal patterns

Seed dispersal by ants (i.e. myrmecochory) is usually considered as a mutualism: ants feed on nutritive bodies, called elaiosomes, before rejecting and dispersing seeds in their nest surroundings. While mechanisms of plant dispersal in the field are well documented, the behaviour of the ant partner was rarely investigated in details. Here, we compared in laboratory conditions the foraging behaviour of two ant species, the omnivorous Lasius niger and the insectivorous Myrmica rubra to which seeds of two European myrmecochorous plants (Chelidonium majus and Viola odorata) were given. Ant colonies were simultaneously presented three types of items: entire seeds with elaiosome (SE), seeds without elaiosome (S) and detached elaiosomes (E). The presence of elaiosomes on seeds did not attract workers from a distance since ants first contact equally each type of items. Although ants are mass-recruiting species, we never observed any recruitment nor trail-laying behaviour towards seeds. For ants having contacted seed items, their antennation, manipulation and seed retrieval behaviour strongly varied depending on the species of each partner. Antennation behaviour, followed by a loss of contact, was the most frequent ant-seed interaction and can be considered as a “hesitation” clue. For both plant species, insectivorous Myrmica ants removed items in larger number and at higher speed than Lasius. This fits with the hypothesis of a convergence between odours of elaiosomes and insect preys. For both ant species, the small Chelidonium seeds were retrieved in higher proportion than Viola ones, confirming the hypothesis that ants prefer the higher elaiosome/diaspore-ratio. Thus, in these crossed experiments, the ant-plant pair Myrmica/Chelidonium was the most effective as ants removed quickly almost all items after a few antennations. The presence of an elaiosome body increased the seed removal by ants excepting for Myrmica which retrieved all Chelidonium seeds, even those deprived of their elaiosome. After 24 h, all the retrieved seeds were rejected out of the nest to the refuse piles. In at least half of these rejected items, the elaiosome was discarded by ants. Species-specific patterns and behavioural differences in the dynamics of myrmecochory are discussed at the light of ant ecology. Received 10 September 2007; revised 5 February 2008; accepted 5 March 2008.     

Small-scale disturbances spread along trophic chains: leaf-cutting ant nests, plants, aphids, and tending ants

Small-scale disturbances caused by animals often modify soil resource availability and may also affect plant attributes. Changes in the phenotype of plants growing on disturbed, nutrient-enriched microsites may influence the distribution and abundance of associated insects. We evaluated how the high nutrient availability generated by leaf-cutting ant nests in a Patagonian desert steppe may spread along the trophic chain, affecting the phenotype of two thistle species, the abundance of a specialist aphid and the composition of the associated assemblage of tending ants. Plants of the thistle species Carduus nutans and Onopordum acanthium growing in piles of waste material generated by leaf-cutting ant nests (i.e., refuse dumps) had more leaves, inflorescences and higher foliar nitrogen content than those in non-nest soils. Overall, plants in refuse dumps showed higher abundance of aphids than plants in non-nest soils, and aphid colonies were of greater size on O. acanthium plants than on C. nutans plants. However, only C. nutans plants showed an increase in aphid abundance when growing on refuse dumps. This resulted in a similar aphid load in both thistle species when growing on refuse dumps. Accordingly, only C. nutans showed an increase in the number of ant species attending aphids when growing on refuse dumps. The increase of soil fertility generated by leaf-cutting ant nests can affect aphid abundance and their tending ant assemblage through its effect on plant size and quality. However, the propagation of small-scale soil disturbances through the trophic chain may depend on the identity of the species involved.     

Increase in ant density promotes dual effects on bee behaviour and plant reproductive performance

Floral rewards do not only attract pollinators, but also herbivores and their predators. Ants are attracted by extrafloral nectaries (EFNs), situated near flowers, and may interfere with the efficiency and behaviour of pollinators. We tested the hypothesis that the impacts of ant–pollinator interactions in plant–pollinator systems are dependent on (1) the seasonal activity of EFNs, which increase ant abundance closer to flowers; (2) consequently, an ant effect, where ants decrease the temporal niche overlap of bees due to predator avoidance; and (3) ant density, where higher densities may negatively affect plant–pollinator interactions and plant performance. We studied two ant–plant–pollinator systems based on Banisteriopsis campestris and Banisteriopsis malifolia plant species. The periods of high ant abundance coincided with plant species blooming. The presence of ants around flowers reduced the visitation rates of the smaller bees and the temporal niche overlap between bee species was not higher than randomly expected when ants had free access. Additionally, we observed variable ant effects on fruit set and duration of bee visits to both Malpighiaceae species when ant density was experimentally kept constant on branches, especially on B. campestris. Our goal was to show the dual role of ant density effects, especially because the different outcomes are not commonly observed in the same plant species. We believe that reduced temporal niche overlap between floral visitors due to ant presence provides an opportunity for smaller bees to improve compatible pollination behaviour. Additionally, we concluded that ant density had variable effects on floral visitor behaviours and plant reproductive performance.     

Respiratory physiology and water relations of three species of Pogonomyrmex harvester ants (Hymenoptera: Formicidae)

The respiratory physiology and water relations of three harvester ant species (Pogonomyrmex rugosus Emery, P. occidentalis[Cresson] and P. californicus[Buckley]) were examined at three temperatures (15, 25 and 35°C) using a flow-through respirometry system. As intact ants tended to be active during testing, we performed a parallel set of experiments on individuals rendered motionless by decapitation. Both intact and decapitated ants exhibited discontinuous ventilation. Decapitation caused metabolic rate (V˙CO₂) and burst frequency to decrease in all three species. Burst volume either remained constant or increased after removal of the head, though mass-specific V˙CO₂ was unaffected except in P. rugosus. Mass-specific V˙CO₂s of headless harvesters were comparable with published values derived from motionless specimens of other ant species. The mean Q₁₀ for intact ants of all three species was 2.37; for decapitated insects the mean was 2.32. Respiratory water constituted a small (< 5%) fraction of total loss, and we believe that discontinuous ventilation does not act to conserve water in these organisms, although it may serve other functions.     

Extranuptial nectaries in flowers: ants increase the reproductive success of the ant‐plant Miconia tococa (Melastomataceae)

• Although the production of extranuptial nectar is a common strategy of indirect defence against herbivores among tropical plants, the presence of extranuptial nectaries in reproductive structures is rare, especially in ant‐plants. This is because the presence of ants in reproductive organs can generate conflicts between the partners, as ants can inhibit the activity of pollinators or even castrate their host plants. Here we evaluate the hypothesis that the ant‐plant Miconia tococa produces nectar in its petals which attracts ants and affects fruit set.
• Floral buds were analysed using anatomical and histochemical techniques. The frequency and behaviour of floral visitors were recorded in field observations. Finally, an ant exclusion experiment was conducted to evaluate the effect of ant presence on fruit production.
• The petals of M. tococa have a secretory epidermis that produces sugary compounds. Nectar production occurred during the floral bud stage and attracted 17 species of non‐obligate ants (i.e. have a facultative association with ant‐plants). Ants foraged only on floral buds, and thus did not affect the activity of pollinators in the neighbouring open flowers. The presence of ants in the inflorescences increased fruit production by 15%.
• To our knowledge, the production of extranuptial nectar in the reproductive structures of a myrmecophyte is very rare, with few records in the literature. Although studies show conflicts between the partners in the ant–plant interaction, ants that forage on M. tococa floral buds protect the plant against floral herbivores without affecting bee pollination.

     

Recruitment in the harvester ant, Pogonomyrmex occidentalis: effects of experimental removal

We examined the importance of experimental removal of mature colonies on colony recruitment in the western harvester ant Pogonomyrmex occidentalis. To test the common assumption that established colonies suppress the establishment of new colonies we removed all colonies in ten 0.25 ha plots in 1996 and an additional five plots in 1997 and measured new colony recruitment in 1997, 1998, and 1999. Using a blocked, paired plot design we found that removal of colonies increased new colony recruitment in some areas of the site, but not others. Spatial variation in the importance of established colonies to recruitment was consistent across years; blocks in which density dependence was important in one year exhibited density dependent recruitment in following years. We estimated that in the blocks where recruitment was affected by established colonies, they accounted for less than 10% of the mortality of foundress queens. The increase in the number of new recruits (on average two additional new colonies) was considerably less than the number of colonies removed; average colony density in the removal plots was 14 colonies per 0.25 ha plot. The consistent lack of importance of established colonies to recruitment in one block and the relatively small response to colony removal in the other blocks suggests that the number of new colonies in a year may not be equivalent to the number of deaths of established colonies in that year. Space limitation is an important influence on recruitment in P. occidentalis, but the magnitude of the limitation varies spatially.     

Testing non-additive effects of nectar-robbing ants and hummingbird pollination on the reproductive success of a parasitic plant

Nectar robbing may have an indirect negative effect on plant reproduction by discouraging legitimate pollinator species from visiting robbed flowers. In this study, we set up a 2 × 2 factorial design with nectar-robbing ants and hummingbird pollination to test for non-additive effects on fruit set, seed mass, and seed germination of the leafless mistletoe Tristerix aphyllus (Loranthaceae). Even though ants caused conspicuous damage at the base of the floral tubes, nectar availability was reduced by only 8 % in the presence of ants. The green-backed firecrown Sephanoides sephaniodes was insensitive to the presence of ants. Rather, the bird responded to flower number and the presence or the absence of damage, but not to the extent of damage within inflorescences. As hummingbirds were largely insensitive to variation in nectar robbing, the interaction ant × hummingbird had no effect on plant-reproductive success. Thus, the factorial experiment did not provide evidence for indirect negative effects of nectar robbing on plant reproduction. These results suggest that indirect effects of nectar robbers on pollinator behaviour may occur under a more restricted set of conditions than those previously considered. We suggest that the low amount of nectar removed by nectar-robbing ants was insufficient for hummingbirds to avoid robbed flowers, which restricted the potential for non-additive effects.     

Relative fitness of aphids: effects of plant quality and ants

We investigated the response of four species of aphids ( Metopeurum fuscoviride , Brachycaudus cardui , Aphis fabae , and Macrosiphoniella tanacetaria ) on tansy ( Tanacetum vulgare ) to plant quality and attendance by an ant, Lasius niger . The aphids experienced one of four different environments for two consecutive generations. Ant-attendance significantly affected the time needed to reach maximum fecundity only in Me. fuscoviride and plant quality in Me. fuscoviride and B. cardui . Maximum daily fecundity was positively affected by plant quality and the magnitude of the effect was inversely associated with the degree of myrmecophily. Ant-attendance had a positive effect on maximum fecundity only in the obligate myrmecophile, Me. fuscoviride . The intrinsic rate of population increase, r _m, on high quality plants, was lowest for the obligate myrmecophile, intermediate for the facultative myrmecophiles and highest for the unattended species. On high quality plants the fitness of Me. fuscoviride was more adversely affected by the developmental stage of the plant and absence of ants than that of A. fabae or Ma. tanacetaria , which were able to maintain a high relative fitness in all the environments. The implications for aphids experiencing different degrees of ant-attendance and seasonal changes in plant quality are discussed.     

Patch colonization by Trirhabda canadensis (Coleoptera: Chrysomelidae): effects of plant species composition and wind

Summary The goldenrod leaf beetle, Trirhabda canadensis, is known to respond to odors of host and non-host species in the laboratory. Here we report movements of T. canadensis in the field in response to volatile odors from monocultures and polycultures of host plants. Overall, beetles preferentially colonized plots with a higher density of host plants and lower diversity of allelochemicals, but under some wind conditions there were marked exceptions. At high windspeeds, they colonized whichever plot(s) was upwind. At low windspeeds, beetles colonized preferred plots even when they were not upwind. The data suggest that odor dispersion varies in a complex way with windspeed: at low windspeeds beetles received information from a wide are of vegetation and made choices while at high windspeeds information was available only from upwind plot(s).