Importance of elaiosome size to removal of ant-dispersed seeds

Several studies relate removal of diaspores from different ant-dispersed plant species to the size of their claiosomes. This study is the first one to relate intraspecific variation in claiosome size to removal of diaspores by ants. This approach circumvents the problem posed by interspecific variation in chemistry and morphology of elaiosomes. We used the systemHepatica nobilis Mill. (Ranunculaceae) andMyrmica ruginodis Nyl. The elaiosome was the attractive part of the diaspore and its attractiveness decreased quickly after release from the fruit. In experimental runs where ants collected diaspores, the elaiosome size of accepted diaspores was larger than of those rejected and the largest diaspores were collected first. Since size of diaspore, elaiosome and achene were correlated, the diaspores that were removed first had both the largest elaiosome and achene. However, our experiments suggested that elaiosome size was more important to removal than achene size or the elaiosome:achene size ratio. If ant dispersal improves plant fitness, elaiosome size and hence diaspore size would be expected to increase over time. However, such directional selection mediated by the ants is probably counterbalanced by the plant. Seed predators and a negative trade-off between number and size of seeds would, among other factors, select for smaller diaspore size,i.e. counteract the effect of the ants' preference for larger elaiosomes.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

Interaction between a threatened endemic plant (<Emphasis Type="Italic">Anchusa crispa</Emphasis>) and the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Seed dispersal mutualisms are essential to ensure the survival of diverse plant species and communities worldwide. Here, we investigated whether the invasive Argentine ant can replace native ants by fulfilling their functional role in the seed dispersal of the rare and threatened endemic myrmecochorous plant, Anchusa crispa, in Corsica (France). Our study addressed the potential of Linepithema humile to disperse elaiosome-bearing seeds of A. crispa, examining L. humile’s effects on (1) the composition of communities of ants removing seeds, (2) the number of seed removals, (3) seed preference, (4) the distance of seed dispersion, and (5) seed germination. We caught seven native species at the control site, but only the Argentine ant at invaded sites. L humile removed A. crispa seeds in greater numbers than did native ants, respectively 66 and 23%, probably due to their higher worker density. The invader was similar to native ants with respect to distance of seed transport. Finally, rates of seed germination were not significantly different between seeds previously in contact with either Argentine ants or not. Taken all together, these results suggest that the Argentine ant is unlikely to pose a threat to A. crispa population. These results have important implications for the management of this rare and threatened endemic plant and provide an example of non-negative interactions between invasive and native species.     

Indirect effects of alternative food resources in an ant–plant interaction

The seeds of many plant species present a food body that is consumed by animal dispersers. In theory, if the animals are polyphagous, the availability of alternative food resource other than the diaspore itself may influence its dispersal and survival. We used the myrmecochore Helleborus foetidus L. (Ranunculaceae), the seeds of which are attached to a lipid-rich elaiosome that is attractive to ants, as a model system to investigate (1) whether alternative foods that are present along with the plant affect ant foraging behavior and diaspore removal and (2) whether food availability in an ant nest affects seed predation and germination. In a field experiment, artificial diaspore depots were offered together with either sugar, insect corpses, seed, or no food (control). Contrary to the prediction that ants would rather concentrate their foraging effort on the highly rewarding alternative foods only, many workers, attracted by the sugar, switched to the hellebore diaspores, which significantly enhanced removal rate. Results obtained in the laboratory further indicated that the larvae of Aphaenogaster iberica (a major seed disperser) predated more on the H. foetidus embryos when no alternative food was available. This, in turn, slightly reduced seed germination. Overall, these results shed light, for the first time, on the potential indirect effects of alternative resources on the fate of diaspores adapted for ant dispersal.     

Geographic variation for elaiosome–seed size ratio and its allometric relationship in two closely related Corydalis species

Abstract

Background: Elaiosomes serve as a reward for seed-dispersing ants. The size of the elaiosome and the elaiosome–seed mass (ESM) ratio are expected to affect seed dispersal, whilst seed size affects the establishment of seedlings. Elaiosomes and seeds can undergo independent change, thus variation in these traits may arise through heterogeneous selection for seedling establishment and for dispersal. Only a few studies have examined intraspecific variation in these traits.

Aims: The aim of this study was to determine if the ratio of elaiosome–seed mass was different in two co-occurring congeneric plant species (Corydalis intermedia and C. solida). Under the hypothesis that the ESM ratio affected diaspore attractiveness to ants, the allometric relationship between elaiosome and seed mass was used to infer how selection by ants may shape the diaspore.

Methods: Seed and elaiosome mass were measured in fruits collected from central and marginal populations of the two plant species. The allometric relationship between seed and elaiosome mass was obtained using major axis regression. In situ cafeteria experiments estimated the removal rate of diaspores by ants.

Results: Intra- and interspecific variation in diaspore traits were found. The selfing C. intermedia produced heavier seeds and elaiosomes than the outcrossing C. solida. Ants removed more diaspores from C. intermedia where both species were present. A slope larger than one characterised the allometric relationship between elaiosome and seed mass in both species. This slope was twice as steep in the central populations of C. intermedia compared to marginal ones.

Conclusions: Results indicate that in C. intermedia elaiosome mass must increase more than proportionally with increasing seed mass for the diaspore to remain attractive to ants. The direction of interspecific differences suggests that a plant-mating system may affect selection for dispersal.     

Signals Can Trump Rewards in Attracting Seed-Dispersing Ants

Both rewards and signals are important in mutualisms. In myrmecochory, or seed dispersal by ants, the benefits to plants are relatively well studied, but less is known about why ants pick up and move seeds. We examined seed dispersal by the ant Aphaenogaster rudis of four co-occurring species of plants, and tested whether morphology, chemical signaling, or the nutritional quality of fatty seed appendages called elaiosomes influenced dispersal rates. In removal trials, ants quickly collected diaspores (seeds plus elaiosomes) of Asarum canadense, Trillium grandiflorum, and Sanguinaria canadensis, but largely neglected those of T. erectum. This discrepancy was not explained by differences in the bulk cost-benefit ratio, as assessed by the ratio of seed to elaiosome mass. We also provisioned colonies with diaspores from one of these four plant species or no diaspores as a control. Colonies performed best when fed S. canadensis diaspores, worst when fed T. grandiflorum, and intermediately when fed A. canadense, T. erectum, or no diaspores. Thus, the nutritional rewards in elaiosomes affected colony performance, but did not completely predict seed removal. Instead, high levels of oleic acid in T. grandiflorum elaiosomes may explain why ants disperse these diaspores even though they reduce ant colony performance. We show for the first time that different elaiosome-bearing plants provide rewards of different quality to ant colonies, but also that ants appear unable to accurately assess reward quality when encountering seeds. Instead, we suggest that signals can trump rewards as attractants of ants to seeds.     

Seed removal decrease by invasive Argentine ants in a high Nature Value farmland

Seed dispersal by ants is an important ecological process that maintains the structure anddiversity of natural communities, however, it is vulnerable to biological invasions. Argentine ants are one of the worst invasive ant species and cause severe changes in ecosystem processes and native ant biodiversity declines in invaded sites. Here, we studied seed removal by ants combining observations and a cafeteria experiment with seeds of four myrmecochorous plant species (Centaurea sphaerocephala, Rosmarinus officinalis, Silybum marianum, and Ulex australis) in two sites (invaded and uninvaded) located in the Mediterranean Montado ecosystem and classified as High Nature Value farmland (HNV). Significant differences in daily seed removal rates were found between the two study sites. In uninvaded sites, several native ant species were attracted to the seeds, resulting in all seeds being removed rapidly. The majority of seed removal events were carried out by two key seed disperses Pheidole pallidula (71%) and Aphaenogaster iberica (26%) with a clear preference for diaspored with larger and heavier elaiosome (i.e., C. sphaerocephala, S. marianum). By contrast, while the Argentine ant showed some interest (68% of seeds were interacted with), no seed removal events were observed. The extirpation of the local ant fauna by the Argentine ant and its inability to ensure seed dispersal services may lead to the interference and eventually to the collapse of seed dispersal of the four studied myrmecochorous plants in the invaded site in the future. We argue that these discrete but severe consequences of an invasive species on a key ecological process may strongly affect the functioning of the Montado ecosystem.     

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.     

Interference competition by Argentine ants displaces native ants: implications for biotic resistance to invasion

The Argentine ant Linepithema humile (Dolichoderinae) is one of the most widespread invasive ant species in the world. Throughout its introduced range, it is associated with the loss or reduced abundance of native ant species. The mechanisms by which these native species are displaced have received limited attention, particularly in Australia. The role of interference competition in the displacement of native ant species by L. humile was examined in coastal vegetation in central Victoria (southeastern Australia). Foragers from laboratory colonies placed in the field consistently and rapidly displaced the tyrant ant Iridomyrmex bicknelli, the big-headed ant Pheidole sp. 2, and the pony ant Rhytidoponera victoriae from baits. Numerical and behavioural dominance enabled Argentine ants to displace these ants in just 20 min; the abundance of native species at baits declined 3.5–24 fold in direct relation to the rapid increase in L. humile. Most precipitous was the decline of I. bicknelli, even though species in this typically dominant genus have been hypothesized to limit invasion of L. humile in Australia. Interspecific aggression contributed strongly to the competitive success of Argentine ants at baits. Fighting occurred in 50–75% of all observed interactions between Argentine and native ants. This study indicates that Argentine ants recruit rapidly, numerically dominate, and aggressively displace from baits a range of Australian native ant species from different subfamilies and functional groups. Such direct displacement is likely to reduce native biodiversity and indirectly alter food web structure and ecosystem processes within invaded areas. Biotic resistance to Argentine ant invasion from native ants in this coastal community in southeastern Australia is not supported in this study.     

Impacts of the invasive Argentine ant on native ants and other invertebrates in coastal scrub in south‐eastern Australia

Invasive ants threaten native biodiversity and ecosystem function worldwide. Although their principal direct impact is usually the displacement of native ants, they may also affect other invertebrates. The Argentine ant, Linepithema humile (Dolichoderinae), one of the most widespread invasive ant species, has invaded native habitat where it abuts peri‐urban development in coastal Victoria in south‐eastern Australia. Here we infer impacts of the Argentine ant on native ants and other litter and ground‐dwelling invertebrates by comparing their abundance and taxonomic composition in coastal scrub forest either invaded or uninvaded by the Argentine ant. Species composition of native ants at bait stations and extracted from litter differed significantly between Argentine ant‐invaded and uninvaded sites and this was consistent across years. Argentine ants had a strong effect on epigeic ants, which were either displaced or reduced in abundance. The native ant Rhytidoponera victoriae (Ponerinae), numerically dominant at uninvaded sites, was completely absent from sites invaded by the Argentine ant. However, small hypogeic ants, including Solenopsis sp. (Myrmicinae) and Heteroponera imbellis (Heteroponerinae), were little affected. Linepithema humile had no detectable effect upon the abundance and richness of other litter invertebrates. However, invertebrate group composition differed significantly between invaded and uninvaded sites, owing to the varied response of several influential groups (e.g. Collembola and Acarina). Floristics, habitat structure and measured environmental factors did not differ significantly between sites either invaded or uninvaded by Argentine ants, supporting the contention that differences in native ant abundance and species composition are related to invasion. Changes in the native ant community wrought by Argentine ant invasion have important implications for invertebrate communities in southern Australia and may affect key processes, including seed dispersal.     

Odor of achlorophyllous plants’ seeds drives seed‐dispersing ants

Seed dispersal by ants is an important means of migration for plants. Many myrmecochorous plants have specialized appendages in their seeds called elaiosome, which provides nutritional rewards for ants, and enable effective seed dispersal. However, some nonmyrmecochorous seeds without elaiosomes are also dispersed by ant species, suggesting the additional mechanisms other than elaiosomes for seed dispersal by ants. The seeds of the achlorophyllous and myco‐heterotrophic herbaceous plant Monotropastrum humile are very small without elaiosomes; we investigated whether odor of the seeds could mediate seed dispersal by ants. We performed a bioassay using seeds of M. humile and the ant Nylanderia flavipes to demonstrate ant‐mediated seed dispersal. We also analyzed the volatile odors emitted from M. humile seeds and conducted bioassays using dummy seeds coated with seed volatiles. Although elaiosomes were absent from the M. humile seeds, the ants carried the seeds to their nests. They also carried the dummy seeds coated with the seed volatile mixture to the nest and left some dummy seeds inside the nest and discarded the rest of the dummy seeds outside the nest with a bias toward specific locations, which might be conducive to germination. We concluded that, in M. humile seeds, volatile odor mixtures were sufficient to induce seed‐carrying behavior by the ants even without elaiosomes.     

其它可利用食物对小花宽瓣黄堇(Corydalis giraldii Fedde)种子散布的影响

作为蚁播植物种子的重要传播者,蚂蚁不但取食种子上附着的油质体,也喜食其它富含蛋白质、脂类、糖和维生素等的食物,因此环境中其它可利用食物的存在可能会影响蚂蚁对种子的搬运进而影响种子散布,但目前对于这种影响是如何发生的仍不清楚。在野外研究了蚂蚁对小花宽瓣黄堇(Corydalis giraldii Fedde)种子、肉、蜂蜜、苹果、馒头等食物的趋性和偏好程度,以及添加食物后蚂蚁对种子的拜访频率和搬运效率,以揭示其它可利用食物如何影响蚂蚁觅食和取食偏好,进而影响小花宽瓣黄堇种子散布。结果显示,在所诱捕的8种蚂蚁中,玉米毛蚁(Lasius alienus(Foerster))和丝光蚁(Formica fusca Linnaeus)是小花宽瓣黄堇种子的主要搬运者,不同食物诱捕的玉米毛蚁数量无显著性差异(P0.05),但蜂蜜和苹果诱捕的丝光蚁数量均显著大于种子(P0.05)。玉米毛蚁和丝光蚁均为杂食性,在觅食中分别行使群体募集和简单协作性募集。在仅有种子的对照处理中,玉米毛蚁和丝光蚁对种子的拜访频率分别为(38.73±4.57)头和(30.8±2.87)头(40min,n=15),两种蚂蚁对种子的拜访频率差异不显著(P0.05);玉米毛蚁和丝光蚁搬运种子的效率分别为(33.87±4.22)粒和(16.27±3.35)粒(40min,n=15),玉米毛蚁的搬运效率显著高于丝光蚁(P0.05)。与对照相比,添加馒头、苹果和蜂蜜后丝光蚁对种子的拜访频率显著降低(P0.05),分别为(15.6±3.61)头、(9.07±1.4)头和(7.67±1.58)头(40min,n=15);添加苹果和蜂蜜后丝光蚁对种子的搬运效率显著降低(P0.05),分别为(3.47±1.17)粒和(2.87±0.9)粒(40min,n=15);添加不同食物后玉米毛蚁对种子的拜访频率和搬运效率均无显著变化(P0.05)。研究结果表明行使群体募集的玉米毛蚁比行使简单协作募集的丝光蚁有更高的种子搬运效率,添加食物后影响丝光蚁对种子的拜访频率和搬运效率,这说明其它可利用食物对小花宽瓣黄堇种子散布的影响与搬运蚂蚁的种类及其觅食的募集方式有关。研究结果可为进一步研究蚂蚁与植物(种子)间的互利共生关系及其影响因素提供资料。     

Are red imported fire ants facilitators of native seed dispersal?

Invasive ants threaten native communities, in part, through their potential to disrupt mutualisms, yet invasive species may also facilitate native species. The red imported fire ant (Solenopsis invicta) is one of the most conspicuous invasive ants in North America and its high densities, combined with its potential to displace native ants, have led to concerns that it may disrupt ant-plant seed dispersal mutualisms. We examined the potential of fire ants to disperse seeds in the longleaf pine ecosystem by comparing the removal of elaiosome-bearing seeds by fire ants versus native ants. A total of 14 ant species were observed removing seeds, with fire ants responsible for more than half of all removals. While fire ants were the dominant seed remover in this system, they did not remove significantly more seeds than would be expected based on their population density (46% of ground-dwelling ants). Moreover, red imported fire ants were similar to native ants with respect to distance of seed movement and frequency of moving seeds back to the nest. Areas of higher fire ant densities were found to have greater rates of seed removal by ants without a subsequent drop in seed dispersal by native ants, suggesting that fire ant-invaded areas may experience overall higher levels of seed dispersal. Thus, fire ants may actually facilitate dispersal of elaiosome-bearing plant species in the longleaf pine ecosystem.     

Opening myrmecochory’s black box: what happens inside the ant nest?

In the process of seed dispersal by ants (myrmecochory), foragers bring diaspores back to their nest, then eat the elaiosome and usually reject viable seeds outside the nest. Here, we investigate what happens inside the nest, a barely known stage of the myrmecochory process, for two seed species (Viola odorata, Chelidonium majus) dispersed either by the insectivorous ant Myrmica rubra or by the aphid-tending ant Lasius niger. Globally, elaiosome detachment decreased ants’ interest towards seeds and increased their probability of rejecting them. However, we found marked differences in seed management by ants inside the nest. The dynamics of elaiosome detachment were ant- and plant-specific whereas the dynamic of seed rejection were mainly ant-specific. Seeds remained for a shorter period of time inside the nest of the carnivorous ant Myrmica rubra than in Lasius niger nest. Thus, elaiosome detachment and seed rejection were two competing dynamics whose relative efficiency leads to variable outcomes in terms of types of dispersed items and of nutrient benefit to the ants. This is why some seeds remained inside the nest even without an elaiosome, and conversely, some seeds were rejected with an elaiosome still attached. Fresh seeds may be deposited directly in contact with the larvae. However, the dynamics of larvae-seeds contacts were also highly variable among species. This study illustrates the complexity and variability of the ecological network of ant–seed interactions.     

Effects of seed morphology and elaiosome chemical composition on attractiveness of five Trillium species to seed‐dispersing ants

Morphological and chemical attributes of diaspores in myrmecochorous plants have been shown to affect seed dispersal by ants, but the relative importance of these attributes in determining seed attractiveness and dispersal success is poorly understood. We explored whether differences in diaspore morphology, elaiosome fatty acids, or elaiosome phytochemical profiles explain the differential attractiveness of five species in the genus Trillium to eastern North American forest ants. Species were ranked from least to most attractive based on empirically‐derived seed dispersal probabilities in our study system, and we compared diaspore traits to test our hypotheses that more attractive species will have larger diaspores, greater concentrations of elaiosome fatty acids, and distinct elaiosome phytochemistry compared to the less attractive species. Diaspore length, width, mass, and elaiosome length were significantly greater in the more attractive species. Using gas chromatography–mass spectrometry, we found significantly higher concentrations of oleic, linoleic, hexadecenoic, stearic, palmitoleic, and total fatty acids in elaiosomes of the more attractive species. Multivariate assessments revealed that elaiosome phytochemical profiles, identified through liquid chromatography–mass spectrometry, were more homogeneous for the more attractive species. Random forest classification models (RFCM) identified several elaiosome phytochemicals that differed significantly among species. Random forest regression models revealed that some of the compounds identified by RFCM, including methylhistidine (α‐amino acid) and d‐glucarate (carbohydrate), were positively related to seed dispersal probabilities, while others, including salicylate (salicylic acid) and citrulline (L‐α‐amino acid), were negatively related. These results supported our hypotheses that the more attractive species of Trillium—which are geographically widespread compared to their less attractive, endemic congeners—are characterized by larger diaspores, greater concentrations of fatty acids, and distinct elaiosome phytochemistry. Further advances in our understanding of seed dispersal effectiveness in myrmecochorous systems will benefit from a portrayal of dispersal unit chemical and physical traits, and their combined responses to selection pressures.     

Disruption of ant-seed dispersal mutualisms by the invasive Asian needle ant (<Emphasis Type="Italic">Pachycondyla chinensis</Emphasis>)

By disrupting the structure of native ant assemblages, invasive ants can have effects across trophic levels. Most studies to date, however, have focused on the impacts just two species (Linepithema humile and Solenopsis invicta). The impacts of many other invasive ant species on ecological processes in their introduced range are unknown. In this study we tested the hypothesis that the invasive ant Pachycondyla chinensis disrupts ant-seed dispersal mutualisms by displacing native ant species, especially the keystone mutualist Aphaenogaster rudis, while failing to disperse seeds itself. In a paired design we measured the impact of P. chinensis on the native ant-plant seed dispersal mutualism. The number of A. rudis workers was 96% lower in invaded than in intact plots, and the number of seeds removed was 70% lower in these plots. Finally, in invaded plots the abundance of Hexastylis arifolia, a locally abundant myrmecochorous plant, was 50% lower than in plots where P. chinensis was absent. A parsimonious interpretation of our results is that P. chinensis causes precipitous declines in the abundance of A. rudis within invaded communities, thereby disrupting the ant-plant seed dispersal mutualisms and reducing abundances of ant-dispersed plants. In sum, the magnitude of the effects of P. chinensis on seed dispersal is quantitatively similar to that documented for the intensively studied invasive Argentine ant. We suggest that more studies on the impacts of less-studied invasive ant species on seed dispersal mutualisms may increase our knowledge of the effects of these invaders on ecosystem function.     

Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. The mechanisms by which Argentine ants effectively compete against native ant species have been previously addressed in field studies that centered on interference and exploitation competition at baits and mainly examined the colony-level performance of Argentine ants. Detailed behavioral observations explaining the basis for the strong competitive ability of L. humile are comparatively rare. To gain a better understanding of the mechanisms by which Argentine ants displace native ants we examined the aggressive interactions between the Argentine ants and the odorous house ant, Tapinoma sessile in four different aggression assays: (1) worker dyad interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a resource competition assay which focused on competition for food and nesting space. Our results demonstrate a clear disparity between worker-level and colony-level fighting ability of Argentine ants and provide behavioral evidence to explain the superior interference ability of Argentine ants in group assays. Argentine ants experienced mixed success in fighting against odorous house ants in dyad interactions, but gradually gained a numerical advantage in symmetrical group interactions by active cooperation among nestmates. Results of the resource competition assay indicate that Argentine ants recruit rapidly, numerically dominate food and nesting sites, and aggressively displace T. sessile from baits. Taken together, the results of these assays allow us to pinpoint the behavioral mechanisms responsible for the remarkable competitive ability of Argentine ants.     

Steep Decline and Cessation in Seed Dispersal by Myrmica rubra Ants

Myrmecochorous diaspores bear a nutrient-rich appendage, the elaiosome, attractive to ant workers that retrieve them into the nest, detach the elaiosome and reject the seed intact. While this interaction is beneficial for the plant partner by ensuring its seed dispersal, elaiosome consumption has various effects −positive, negative or none − on ants’ demography and survival, depending on both the ant/plant species involved. In this context, the contribution of ants to seed dispersal strongly varies according to the ant/plant pairs considered. In this paper, we investigate whether the dynamics of myrmecochory also vary on a temporal scale, for a given pair of partners: Myrmica rubra ants and Viola odorata seeds. During their first encounter with seeds, ants collect all the diaspores and eat the majority of elaiosomes. Both the harvesting effort and the elaiosome consumption decline when seeds are offered on the next week and completely cease for the following weeks. This is related to a decrease in the number of foragers reaching the food source, as well as to a reduced probability for an ant contacting a seed to retrieve it. Seed retrieval is not reactivated after seven weeks without any encounter with V. odorata seeds. By contrast, naive ant colonies only fed with fruit flies do not show a decline of prey harvesting of which the speed of retrieval even increases over the successive weeks. Myrmecochory may thus be labile at the scale of a fruiting season due to the ability of ants to steeply tune and cease for several months the harvesting of these seemingly poorly rewarding items and to maintain cessation of seed exploitation. The present study emphasizes the importance of a long-lasting follow up of the myrmecochory process, to assess the stability of this ant-plant partnership and to identify mechanisms of adaptive harvesting in ants.     

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.     

Colony-structure variation and interspecific competitive ability in the invasive Argentine ant

The success of some invasive species may depend on phenotypic changes that occur following introduction. In Argentine ants ( Linepithema humile) introduced populations typically lack intraspecific aggression, but native populations display such behavior commonly. We employ three approaches to examine how this behavioral shift might influence interspecific competitive ability. In a laboratory experiment, we reared colonies of Forelius mccooki with pairs of Argentine ant colonies that either did or did not exhibit intraspecific aggression. F. mccooki reared with intraspecifically non-aggressive pairs of Argentine ants produced fewer eggs, foraged less actively, and supported fewer living workers than those reared with intraspecifically aggressive pairs. At natural contact zones between competing colonies of L. humile and F. mccooki, the introduction of experimental Argentine ant colonies that fought with conspecific field colonies caused L. humile to abandon baits in the presence of F. mccooki, whereas the introduction of colonies that did not fight with field colonies of Argentine ants resulted in L. humile retaining possession of baits. Additional evidence for the potential importance of colony- structure variation comes from the Argentine ants native range. At a site along the Rio de la Plata in Argentina, we found an inverse relationship between ant richness and density of L. humile (apparently a function of local differences in colony structure) in two different years of sampling.