Seed handling behaviours of native and invasive seed‐dispersing ants differentially influence seedling emergence in an introduced plant

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Myrmecochory and short-term seed fate in Rhamnus alaternus: Ant species and seed characteristics

Benefits conferred on plants in ant-mediated seed dispersal mutualisms (myrmecochory) depend on the fate of transported seeds. We studied the effects of elaiosome presence, seed size and seed treatment (with and without passage through a bird's digestive tract) on short-term seed fate in Rhamnus alaternus. In our study, we define short-term seed, or initial, seed fate, as the location where ants release the seeds after ant contact with it. The elaiosomes had the most influence on short-term fate, i.e. whether or not seeds were transported to the nest. The workers usually transported big seeds more often than small ones, but small ants did not transport large seeds. Effect of seed size on transport depended on the ant species and on the treatment of the seed (manual extraction simulating a direct fall from the parent plant vs. bird deposition corresponding to preliminary primary dispersal). Probability of removal of elaiosome-bearing seeds to the nest by Aphaenogaster senilis increased with increasing seed weight.     

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.     

New mutualism for old: indirect disruption and direct facilitation of seed dispersal following Argentine ant invasion

The indirect effects of biological invasions on native communities are poorly understood. Disruption of native ant communities following invasion by the Argentine ant (Linepithema humile) is widely reported to lead indirectly to the near complete collapse of seed dispersal services. In coastal scrub in southeastern Australia, we examined seed dispersal and handling of two native and two invasive alien plant species at Argentine ant-invaded or -uninvaded sites. The Argentine ant virtually eliminates the native keystone disperser Rhytidoponera victoriae, but seed dispersal did not collapse following invasion. Indeed, Argentine ants directly accounted for 92% of all ant-seed interactions and sustained overall seed dispersal rates. Nevertheless, dispersal quantity and quality among seed species differed between Argentine ant-invaded and -uninvaded sites. Argentine ants removed significantly fewer native Acacia retinodes seeds, but significantly more small seeds of invasive Polygala myrtifolia than did native ants at uninvaded sites. They also handled significantly more large seeds of A. sophorae, but rarely moved them >5 cm, instead recruiting en masse, consuming elaiosomes piecemeal and burying seeds in situ. In contrast, Argentine ants transported and interred P. myrtifolia seeds in their shallow nests. Experiments with artificial diaspores that varied in diaspore and elaiosome masses, but kept seed morphology and elaiosome quality constant, showed that removal by L. humile depended on the interaction of seed size and percentage elaiosome reward. Small diaspores were frequently taken, independent of high or low elaiosome reward, but large artificial diaspores with high reward instead elicited mass recruitment by Argentine ants and were rarely moved. Thus, Argentine ants appear to favour some diaspore types and reject others based largely on diaspore size and percentage reward. Such variability in response indirectly reduces native seed dispersal and can directly facilitate the spread of an invasive alien shrub.     

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.     

蚂蚁觅食与搬运行为对阜平黄堇和小花黄堇种子散布的影响

蚂蚁是无脊椎动物中重要的种子传播者,蚂蚁散布影响植物种子的传播和扩散,进而会影响种苗的空间分布格局。在野外研究了蚂蚁觅食及搬运行为对阜平黄堇(Corydalis wilfordii Regel)和小花黄堇(C.racemosa(Thunb.)Pers.)种子散布的影响。结果显示,双针棱胸蚁和束胸平结蚁是两种植物种子的共同搬运者,前者行使群体募集,后者行使简单协作募集。在搬运阜平黄堇种子时,双针棱胸蚁在原地或搬运途中取食油质体后抛弃的种子约占种子总数的56%,而拖至蚁巢的种子约占种子总数的44%,平均搬运距离为(1.85±0.24)m,搬运效率为(43.8±7.5)粒/h;而束胸平结蚁将完整种子全部直接搬运至蚁巢,平均搬运距离为0.45 m,搬运效率为(7.3±2.2)粒/h。在搬运小花黄堇种子时,双针棱胸蚁和束胸平结蚁均将完整种子全部直接搬运至蚁巢,平均搬运距离分别为(6.27±4.40)m和(6.65±1.64)m,搬运效率分别为(34.2±6.5)粒/h和(10.6±3.2)粒/h。这说明行使群体募集的蚂蚁比行使简单协作募集的蚂蚁有较高的搬运效率,蚂蚁散布导致阜平黄堇和小花黄堇种子到达蚁巢的数量和搬运距离不同,而这种不同与相应搬运蚂蚁的觅食对策、搬运行为和种子特征有关。阜平黄堇种子比小花黄堇种子大,但阜平黄堇的油质体质量比小于小花黄堇的油质体质量比,讨论了种子特征对蚂蚁散布的影响。     

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.     

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.     

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.     

Harvester ant seed removal in an invaded sagebrush ecosystem: Implications for restoration

A better understanding of seed movement in plant community dynamics is needed, especially in light of disturbance‐driven changes and investments into restoring degraded plant communities. A primary agent of change within the sagebrush‐steppe is wildfire and invasion by non‐native forbs and grasses, primarily cheatgrass (Bromus tectorum). Our objectives were to quantify seed removal and evaluate ecological factors influencing seed removal within degraded sagebrush‐steppe by granivorous Owyhee harvester ants (Pogonomyrmex salinus Olsen). In 2014, we sampled 76 harvester ant nests across 11 plots spanning a gradient of cheatgrass invasion (40%–91% cover) in southwestern Idaho, United States. We presented seeds from four plant species commonly used in postfire restoration at 1.5 and 3.0 m from each nest to quantify seed removal. We evaluated seed selection for presented species, monthly removal, and whether biotic and abiotic factors (e.g., distance to nearest nest, temperature) influenced seed removal. Our top model indicated seed removal was positively correlated with nest height, an indicator of colony size. Distance to seeds and cheatgrass canopy cover reduced seed removal, likely due to increased search and handling time. Harvester ants were selective, removing Indian ricegrass (Achnatherum hymenoides) more than any other species presented. We suspect this was due to ease of seed handling and low weight variability. Nest density influenced monthly seed removal, as we estimated monthly removal of 1,890 seeds for 0.25 ha plots with 1 nest and 29,850 seeds for plots with 15 nests. Applying monthly seed removal to historical restoration treatments across the western United States showed harvester ants can greatly reduce seed availability at degraded sagebrush sites; for instance, fourwing saltbush (Atriplex canescens) seeds could be removed in <2 months. Collectively, these results shed light on seed removal by harvester ants and emphasize their potential influence on postfire restoration within invaded sagebrush communities.     

Seed predator deterrence by seed-carrying ants in a dyszoochorous plant, <Emphasis Type="Italic">Chamaesyce maculata</Emphasis> L. Small (Euphorbiaceae)

Seeds are often carried by omnivorous ants even if they do not carry elaiosomes. Although many seeds carried by ants are consumed, both seeds abandoned during the seed carrying and leftover seeds are consequently dispersed (dyszoochory). These non-myrmecochorous seeds do not necessarily attract ants quickly. Therefore, these seeds often seem to be exposed to the danger of consumption by pre-dispersal seed predators. We propose the hypotheses, “seed predator deterrence hypothesis” that plants may benefit from seed-carrying ants if they deter seed predators from visiting plants, and seed-carrying ants may play additional roles in plant reproductive success, besides dyszoochory by ants. To test the hypotheses, we investigated the abundance of seed-carrying ants of the species Tetramorium tsushimae Linnaeus and Pheidole noda Smith F., and of the seed predatory stinkbug, Nysius plebeius Distat, on the spotted sandmat, Chamaesyce maculata L. Small, of which the seeds have no elaiosomes but are consumed by both ants and bugs. In the field, ants and stinkbugs seldom encountered each other on the plant. The number of stinkbugs beneath the plants with ants was smaller than that beneath the plants without ants. In laboratory experiments, the number of stinkbugs on the shoot was smaller when ants were present than when they were absent. These results might support the seed predator deterrence hypothesis: the probability of seed predation by stinkbugs seems to be reduced by the ant visits on plants and/or the existence of ants beneath the plants. This study highlights a new ant–plant interaction in seed dispersal by ants.     

DUAL FUNCTION OF THE ELAIOSOME OF CORYDALIS AUREA (FUMARIACEAE): ATTRACTION OF DISPERSAL AGENTS AND REPULSION OF PEROMYSCUS MANICULATUS,A SEED PREDATOR

It has been suggested that one of the selective advantages of ant dispersal is the burial of seeds in ant nests where predators such as small rodents cannot find them. The elaiosomes of Corydalis aurea (Fumariaceae) are extremely attractive to ants, which assiduously gather the seeds and take them to nests. However, seed production commonly exceeds the gathering capacity of ants so that seeds accumulate beneath the parent plants. In spite of this, no signs of rodent predation are evident. Experiments with a major seed predator, the deer mouse Peromyscus maniculatus, show that when given the choice of seeds with and without elaiosomes, the mice consume significantly more seeds without elaiosomes. This remains true whether or not the intact seeds bear fresh, moist elaiosomes or dry, withered ones. Our experiments strongly suggest that the elaiosome has a dual function, the attraction of the ant seed dispersers and the repulsion of seed predators.     

DISPERSAL ECOLOGY OF CAREX PEDUNCULATA (CYPERACEAE), A NEW NORTH AMERICAN MYRMECOCHORE

Carex pedunculata is the first North American species of the Cyperaceae that is identified as a myrmecochore. Many morphological and phenological features of this species and its breeding system are interpreted as adaptive for seed dispersal by ants. In laboratory tests, workers of the ant species Aphaenogaster rudis carry the diaspores to the nest, eat the elaiosomes, carry larvae to the elaiosomes to feed, and deposit diaspores whose elaiosomes have been eaten with other nest debris. The achenes then germinate. Achenes will also germinate without any handling by ants. Workers will also transport diaspores with uneaten elaiosomes when the nest is disturbed. Greenhouse tests show that seedling growth is greatly inhibited if a diaspore remains near the parent plant and cohort seedlings. Field studies of natural populations identify rotting logs (the location of ant nests) as forest floor microsites for colonization of C. pedunculata and other myrmecochores. Ant nesting behavior may pattern much of the herb stratum. This species is self-compatible, and single seeds may start successful new populations. Three processes contribute to population growth: vegetative growth, germination of untransported diaspores, and germination of ant-transported diaspores.     

Differential dispersal of Chamaesyce maculata seeds by two ant species in Japan

Seed dispersal by ants (myrmecochory) is a widely distributed plant–animal interaction in many ecosystems, and it has been regarded as a generalized (multiple species) interaction in which specialization on specific ant partners is uncommon. In this paper, we demonstrate species-specific seed dispersal of spotted spurge (Chamaesyce maculata) by ants in Japan. C. maculata produces seeds from summer to autumn in Japan. The seeds produced in autumn are carried by two ant species, Tetramorium tsushimae and Pheidole noda. We performed laboratory experiments to investigate the fate of C. maculata seeds in the nests of T. tsushimae and P. noda. P. noda consumed the seeds in the nest and rarely carried seeds out of the nest, while T. tsushimae consumed only the seed coat, and subsequently carried the seeds out of the nest. Removal of the seed coat by T. tsushimae may increase seed survival by reducing their susceptibility to infection by fungi. We also observed ant responses to filter paper soaked with an aqueous extract of the seed coat. P. noda did not react to the filter paper, but T. tsushimae carried the filter paper into their nest. Analysis by high-pressure liquid chromatography revealed that the aqueous extract contained at least four sugars and one unknown substance. Myrmecochory has been regarded as a generalized interaction with specialization for specific ant partners uncommon. However, our study suggests there is a species-specific interaction in seed dispersal by ants in autumn-flowering individuals of C. maculata in Japan.     

Effects of ant behaviour and presence of extrafloral nectaries on seed dispersal of the Neotropical myrmecochore Turnera ulmifolia L. (Turneraceae)

This paper describes the myrmecochory system of Turnera ulmifolia in a coastal sand dune matorral in Mexico. Turnera ulmifolia has elaiosome‐bearing seeds and extrafloral nectaries (EFNs). In ten quadrants (4 × 15 m) ant–seed interaction was monitored, and an interaction intensity index calculated and correlated with the number of seedlings. Seed removal rates by ants were surveyed every 2 h for 24 h, the ants being observed both on and beneath the plants. The role of the elaiosome in seed removal was evaluated by offering seeds with and without elaiosomes, and elaiosomes only. Finally, the effect of ant manipulation in seed germination was evaluated. There were 25 ant species associated with seeds and/or EFNs, the most frequently recorded being Monomorium cyaneum and Forelius analis. There was a positive correlation between the intensity index and seedling number per quadrant. There was significantly higher mean seed removal during the day than during the night (19.3% and 12.3%, respectively), and from beneath than on the plant (21.9% and 9.5%, respectively). The preference for elaiosomes only was also greater during the diurnal period, and when gathered on, rather than beneath, the plant. Seed manipulation by F. analis enhanced germination by T. ulmifolia. Seed removal, dispersal distances, seed predation and germination were largely determined by ant behaviour. The presence of EFNs may be influencing seed removal on the plant by attracting a specific assemblage of omnivorous ants. Among such assemblages associated with T. ulmifolia we encountered a variety of behaviours, with ant species either good at defending plants but bad at dispersing seeds, or vice versa. We discuss the way in which these two rewards, and the processes involved (defence and dispersion), could have interacted with each other and evolved. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 67–77.     

Multiphase myrmecochory: the roles of different ant species and effects of fire

Seed dispersal by ants (myrmecochory) can be influenced by changes to ant assemblages resulting from habitat disturbance as well as by differences in disperser behaviour. We investigated the effect of habitat disturbance by fire on the dispersal of seeds of a myrmecochorous shrub, Pultenaea daphnoides. We also investigated the consequence of the seed relocation behaviours of two common dispersers (Pheidole sp. A and Rhytidoponera metallica) for the redispersal of seeds. Pheidole sp. A colonies did not relocate seeds outside their nests. In contrast, R. metallica colonies relocated 43.6 % of seeds fed to them, of which 96.9 % had residual elaiosome that remained attached. On average, R. metallica relocated seeds 78.9 and 60.7 cm from the nest entrances in burned and unburned habitat, respectively. Seeds were removed faster in burned than in unburned habitat, and seeds previously relocated by R. metallica were removed at similar rates to seeds with intact elaiosomes, but faster than seeds with detached elaiosomes. Dispersal distances were not significantly different between burned (51.3 cm) and unburned (70.9 cm) habitat or between seeds with different elaiosome conditions. Differences between habitat types in the frequency of seed removal, the shape of the seed dispersal curve, and the relative contribution of R. metallica and Pheidole sp. A to seed dispersal were largely due to the effect of recent fire on the abundance of Pheidole sp. A. Across habitat types, the number of seeds removed from depots and during dispersal trials most strongly related to the combined abundances of R. metallica and Pheidole. Our findings show that myrmecochory can involve more than one dispersal phase and that fire indirectly influences myrmecochory by altering the abundances of seed-dispersing ants.     

Differences in reward removal efficiency and its consequences for seed germination in the ant-dispersed sedge <Emphasis Type="Italic">Carex tristachya</Emphasis> (Cyperaceae)

Reward removal is an essential step for seed dispersal mutualism because residual rewards inhibit germination. Nevertheless, variation in the reward removal efficiency (RRE) among dispersers and its consequences for germination have rarely been reported. In this study, we compared the RREs of two sympatric seed-dispersing ants, Formica japonica and Pheidole noda, using seeds of the ant-dispersed sedge Carex tristachya. Then, we conducted seed sowing experiments in a non-heated glasshouse to evaluate the effect of RRE on the percentage and speed of germination. The majority (85%) of seeds handled by F. japonica had residual elaiosomes, while elaiosomes were completely removed from all seeds handled by P. noda, demonstrating that P. noda has much higher RRE than F. japonica. The seed sowing experiments revealed that RRE, defined by the presence or absence of residual elaiosomes, was not associated with the percentage germination within a year. However, high RRE seeds with no residual elaiosomes germinated significantly faster than low RRE seeds with residual elaiosomes. Similarly, artificial removal of elaiosomes from C. tristachya seeds accelerated germination speed without affecting germination percentage. These results suggest that RRE is one of the most important parameters determining the effectiveness of a seed dispersal agent.     

SEED DISPERSAL OF TRILLIUM OVATUM (LILIACEAE) IN SECOND-GROWTH REDWOOD FORESTS

Trillium ovatum (Liliaceae) is myrmecochorous: its seeds bear large elaiosomes that are attractive to ants. Nevertheless, in coastal second-growth redwood forests of northern California, most seedlings occur in mixed-age clusters close to potential parents, suggesting that seed dispersal is limited. Ants were absent or rare at two relatively cool, moist study sites. At these sites, most seeds either eventually fell passively from fruits or were knocked to the ground by banana slugs that foraged on the elaiosomes. At two warmer, drier sites, a single species of ant, Lasius pallitarsis, dispersed the seeds but tended to remove the elaiosomes before returning to the nest. Thus at all sites a large number of seeds remained close to adults, accounting for the observed pattern of seedling distribution. The dispersal adaptations of T. ovatum and other redwood forest myrmecochores probably evolved in forests where seed-carrying ants were more common.     

Dropping rates of elaiosome-bearing seeds during transport by ants (Formica polyctena Foerst.): Implications for distance dispersal

In a deciduous forest, foraging ants collect elaiosome-bearing seeds and carry them to their nests. Some of the seeds reach the nest and are concentrated there. Others may be dropped by ants during transport. The dropped seeds enter the soil seed pool. However, they might be repeatedly removed by other ant individuals and carried again in the direction of the nest. Rates of seed dropping and repeated removals must be known to evaluate the effect of ant workers on dispersal distance of seeds. The rate of seed dropping is predicted to depend on size of seeds and of elaiosomes, both of which vary among plant species, and on the size of the ant workers. Mark-recapture experiments were used to evaluate dropping rates of seeds of five myrmecochorous and diplochorous plants (Chelidonium majus L., Asarum europaeum L., Viola matutina Klok., V. mirabilis L., V. hirta L.) during their transport by the ant Formica polyctena Foerst. In the series of species A. europaeum – V. hirta – V. mirabilis – Ch. majus – V. matutina, the dropping rate increased. Small workers dropped seeds of A. europaeum more often than did large ones, while seeds of V. hirta were dropped by ants of different size classes with the same frequency. Across species, dropping rates of seeds were negatively correlated with the rate at which ants removed them from the depot. The number of seeds which reach the nests is the other important parameter of seed dispersal. This parameter depends on dropping rates: seeds with lower dropping rates have higher chances of being deposited in nests. Diplochores usually produce many small seeds, which are characterised by low removal rates and high dropping rates during transport by ants. Obligate myrmecochores produce rather few large seeds, which have high removal rates and low dropping rates. To analyse the significance of seed dropping in the dispersal distance of seeds, a computer simulation based on two factors [(i) seed number produced by a plant; (ii) dropping rate of seeds] is proposed.     

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.