Species-Specific Seed Dispersal in an Obligate Ant-Plant Mutualism

Throughout lowland Amazonia, arboreal ants collect seeds of specific plants and cultivate them in nutrient-rich nests, forming diverse yet obligate and species-specific symbioses called Neotropical ant-gardens (AGs). The ants depend on their symbiotic plants for nest stability, and the plants depend on AGs for substrate and nutrients. Although the AGs are limited to specific participants, it is unknown at what stage specificity arises, and seed fate pathways in AG epiphytes are undocumented. Here we examine the specificity of the ant-seed interaction by comparing the ant community observed at general food baits to ants attracted to and removing seeds of the AG plant Peperomia macrostachya. We also compare seed removal rates under treatments that excluded vertebrates, arthropods, or both. In the bait study, only three of 70 ant species collected P. macrostachya seeds, and 84% of observed seed removal by ants was attributed to the AG ant Camponotus femoratus. In the exclusion experiment, arthropod exclusion significantly reduced seed removal rates, but vertebrate exclusion did not. We provide the most extensive empirical evidence of species specificity in the AG mutualism and begin to quantify factors that affect seed fate in order to understand conditions that favor its departure from the typical diffuse model of plant-animal mutualism.     

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.     

First report of seed dispersal by ants in Dicentra peregrina (Papaveraceae), an alpine plant in the Japanese Alps

Seed dispersal by ants, known as myrmecochory, is commonly observed among various plant taxa. The seeds of these plants have an elaiosome to attract ants. In Japan, myrmecochory has been well studied in several lowland plant species, but not in highland plant species that grow above the tree line. We investigated whether the seeds of Dicentra peregrina, known as the “queen of the alpine plants” in Japan, are carried by Formica gagatoides ants at 2510 m a.s.l. on Mt Norikura, Kita‐Alps, Japan. We observed F. gagatoides workers picking up D. peregrina seeds by grasping the elaiosome and carrying the seeds into their nests. We inferred from the observed ant behavior and the seed morphology that D. peregrina is a myrmecochorous species.     

Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests

Summary Of 36 plant species surveyed, 6 were significantly associated with nests of the desert seed-harvester ant Veromessor pergandei or Pogonomyrmex rugosus; two other plant species were significantly absent from ant nests. Seeds of two common desert annuals, Schismus arabicus and Plantago insularis, realize a 15.6 and 6.5 fold increase (respectively) in number of fruits or seeds produced per plant growing in ant nest refuse piles compared to nearby controls. Mass of individual S. arabicus seed produced by plants growing in refuse piles also increased significantly. Schismus arabicus, P. insularis and other plants associated with ant nests do not have seeds with obvious appendages attractive to ants. Dispersal and reproductive increase of such seeds may represent a relatively primitive form of ant-plant dispersal devoid of seed morphological specializations. Alternatively, evolution of specialized seed structures for dispersal may be precluded by the assemblage of North American seed-harvester ants whose workers are significantly larger than those ants normally associated with elaiosome-attached seed dispersal. Large worker size may permit consumption of elaiosome and seed.     

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。     

Seed dispersal by ants in the semi-arid Caatinga of North-East Brazil

BACKGROUND AND AIMS: Myrmecochory is a conspicuous feature of several sclerophyll ecosystems around the world but it has received little attention in the semi-arid areas of South America. This study addresses the importance of seed dispersal by ants in a 2500-km(2) area of the Caatinga ecosystem (north-east Brazil) and investigates ant-derived benefits to the plant through myrmecochory. METHODS: Seed manipulation and dispersal by ants was investigated during a 3-year period in the Xingó region. Both plant and ant assemblages involved in seed dispersal were described and ant behaviour was characterized. True myrmecochorous seeds of seven Euphorbiaceae species (i.e. elaiosome-bearing seeds) were used in experiments designed to: (1) quantify the rates of seed cleaning/removal and the influence of both seed size and elaiosome presence on seed removal; (2) identify the fate of seeds dispersed by ants; and (3) document the benefits of seed dispersal by ants in terms of seed germination and seedling growth. KEY RESULTS: Seed dispersal by ants involved one-quarter of the woody flora inhabiting the Xingó region, but true myrmecochory was restricted to 12.8 % of the woody plant species. Myrmecochorous seeds manipulated by ants faced high levels of seed removal (38-84 %) and 83 % of removed seeds were discarded on ant nests. Moreover, seed removal positively correlated with the presence of elaiosome, and elaiosome removal increased germination success by at least 30 %. Finally, some Euphorbiaceae species presented both increased germination and seedling growth on ant-nest soils. CONCLUSIONS: Myrmecochory is a relevant seed dispersal mode in the Caatinga ecosystem, and is particularly frequent among Euphorbiaceae trees and shrubs. The fact that seeds reach micro-sites suitable for establishment (ant nests) supports the directed dispersal hypothesis as a possible force favouring myrmecochory in this ecosystem. Ecosystems with a high frequency of myrmecochorous plants appear not to be restricted to regions of nutrient-impoverished soil or to fire-prone regions.     

Differential Recruitment of Camponotus femoratus (Fabricius) Ants in Response to Ant Garden Herbivory

Although several studies have shown that ants can recognize chemical cues from their host plants in ant-plant systems, it is poorly demonstrated in ant gardens (AGs). In this interaction, ant species constantly interact with various epiphyte species. Therefore, it is possible to expect a convergence of chemical signals released by plants that could be acting to ensure that ants are able to recognize and defend epiphyte species frequently associated with AGs. In this study, it was hypothesized that ants recognize and differentiate among chemical stimuli released by AG epiphytes and non-AG epiphytes. We experimentally simulated leaf herbivore damage on three epiphyte species restricted to AGs and a locally abundant understory herb, Piper hispidum, in order to quantify the number of recruited Camponotus femoratus (Fabricius) defenders. When exposed to the AG epiphytes Peperomia macrostachya and Codonanthe uleana leaves, it was observed that the recruitment of C. femoratus workers was, on average, respectively 556% and 246% higher than control. However, the number of ants recruited by the AG epiphyte Markea longiflora or by the non-AG plant did not differ from paper pieces. This indicated that ants could discern between chemicals released by different plants, suggesting that ants can select better plants. These results can be explained by evolutionary process acting on both ants’ capability in discerning plants’ chemical compounds (innate attraction) or by ants’ learning based on the epiphyte frequency in AGs (individual experience). To disentangle an innate behavior, a product of classical coevolutionary process, from an ant’s learned behavior, is a complicated but important subject to understand in the evolution of ant-plant mutualisms.     

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

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Mutualism fails when climate response differs between interacting species

Successful species interactions require that both partners share a similar cue. For many species, spring warming acts as a shared signal to synchronize mutualist behaviors. Spring flowering plants and the ants that disperse their seeds respond to warming temperatures so that ants forage when plants drop seeds. However, where warm‐adapted ants replace cold‐adapted ants, changes in this timing might leave early seeds stranded without a disperser. We investigate plant seed dispersal south and north of a distinct boundary between warm‐ and cold‐adapted ants to determine if changes in the ant species influence local plant dispersal. The warm‐adapted ants forage much later than the cold‐adapted ants, and so we first assess natural populations of early and late blooming plants. We then transplant these plants south and north of the ant boundary to test whether distinct ant climate requirements disrupt the ant–plant mutualism. Whereas the early blooming plant's inability to synchronize with the warm‐adapted ant leaves its populations clumped and patchy and its seedlings clustered around the parents in natural populations, when transplanted into the range of the cold‐adapted ant, effective seed dispersal recovers. In contrast, the mutualism persists for the later blooming plant regardless of location because it sets seed later in spring when both warm‐ and cold‐adapted ant species forage, resulting in effective seed dispersal. These results indicate that the climate response of species interactions, not just the species themselves, is integral in understanding ecological responses to a changing climate. Data linking phenological synchrony and dispersal are rare, and these results suggest a viable mechanism by which a species' range is limited more by biotic than abiotic interactions – despite the general assumption that biotic influences are buried within larger climate drivers. These results show that biotic partner can be as fundamental a niche requirement as abiotic resources.     

The effects of harvester ant (Messor ebeninus Forel) nests on vegetation and soil properties in a desert dwarf shrub community in north-eastern Arabia

In desert ecosystems, harvester ants have been shown to be important granivores and seed dispersers. Because many desert plants exist as seeds in the ground for long periods, harvester ants may greatly influence plant population dynamics. In this study, we examine the effects of harvester ant nests on vegetation and soil properties in a desert ecosystem dominated by the dwarf shrub Rhanterium epapposum in Kuwait, north-eastern Arabia. Soil properties were greatly modified in the circular refuse zone around the nests of the harvester ants, with elevated levels of nutrients (except nitrogen) and organic matter. Plant species richness and productivity were also significantly enhanced in the refuse zone. The vast majority of species inhabiting this zone were desert annuals, but there is no evidence to suggest that the species involved have any distinct association with ant nests. Harvester ants therefore contribute substantially to small-scale spatial heterogeneity in this aridland plant community. It is also suggested that the significance of ant nests in providing favourable sites for plant growth is enhanced in drier years, when many species in off-nest sites may die prematurely due to drought. Furthermore, because ants occasionally abandon their nests and create new colonies, the addition of nutrient-rich patches to the landscape over time could represent an important mechanism for maintaining fertility of desert soils, possibly with long-term implications for plant biodiversity. Harvester ants can therefore be regarded as a key ecosystem engineer in this relatively undisturbed desert ecosystem of Kuwait.     

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.     

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.     

Secondary seed dispersal by ants in Neotropical cerrado savanna: species‐specific effects on seeds and seedlings of Siparuna guianensis (Siparunaceae)

1. Most woody plant species in tropical habitats are primarily vertebrate‐dispersed, but interactions between ants and fallen seeds and fruits are frequent. This study assesses the species‐specific services provided by ants to fallen arillate seeds of Siparuna guianensis, a primarily bird‐dispersed tree in cerrado savanna. The questions of which species interact with fallen seeds, their relative contribution (versus vertebrates) to seed removal, and the potential effects on seedling establishment are investigated. 2. Seeds are removed in similar quantities in caged and control treatments, suggesting that ants are the main dispersers on the ground. Five ant species attended seeds. Pheidole megacephala (≈0.4 cm) cooperatively transported seeds, whereas the smaller Pheidole sp. removed the seed aril on spot. Large (> 1.0 cm) Odontomachus chelifer, Pachycondyla striata, and Ectatomma edentatum individually carried seeds up to 4 m. Bits of aril are fed to larvae and intact seeds are discarded near the nest entrance. 3. Overall, greater numbers of seedlings were recorded near ant nests than in control plots without nests. This effect, however, was only detected near P. megacephala and P. striata nests, where soil penetrability was greater compared with controls. Soil nutrients did not differ between paired plots. 4. This study confirms the prevalence of ant–seed interactions in cerrado and shows that ant‐derived benefits are species‐specific. Ant services range from seed cleaning on the spot to seed displacement promoting non‐random spatial seedling recruitment. Although seed dispersal distances by ants are likely to be shorter than those by birds, our study of S. guianensis shows that fine‐scale ant‐induced seed movements may ultimately enhance plant regeneration in cerrado.     

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.     

Mucilage secretion by seeds doubles the chance to escape removal by ants

Post-dispersal seed predation is a risk for plants in semiarid environments, leading to strategies to protect their propagules from seed collection by animals. In this study, we evaluated the importance of mucilage secretion in seeds as a mechanism to reduce seed collection by ants. We selected three Mediterranean species with strong mucilage secretion on their seeds which become sticky upon wetting. Seeds of Rosmarinus officinalis, Fumana ericoides, and Fumana thymifolia were exposed to ants and survival was compared between dry loose seeds and seeds glued to the soil with previously secreted mucilage. The study site was in the Sierra Calderona, 25?km north of Valencia (Spain). The ant?Cplant interaction was analyzed by scrutinizing seed collection by ants and by analyzing the waste piles of ant nests. To test survival, groups of 10 seeds were placed on the ground. Each group consisted of five control (dry) and five?mucilaginous seeds (previously mucilage secreted) and was covered by the cover of a Petri dish modified to permit only the entry of ants. Seeds were inspected weekly for seed disappearance and the survival function (Kaplan?CMeier estimator) was calculated. Seeds of the target species were important food items for ants and were actively collected, and more than 50?% of the experimental seeds that were glued to the ground with their own mucilage survived at the end of the study period but only 0?C20?% of the control seeds survived after the same time of exposure. The implications for plant establishment of these findings are discussed.     

Switching roles from antagonist to mutualist: a harvester ant as a key seed disperser of a myrmecochorous plant

1. Harvester ants are major seed predators in arid environments. However, given that many harvester ants are partly omnivorous and therefore potentially attracted to the elaiosomes of myrmecochorous seeds, it is unclear if these ants act as predators or dispersers when removing myrmecochorous seeds. 2. We describe the outcomes of interactions between the harvester ant, Pogonomyrmex naegelii, and myrmecochorous plant, Microstachys serrulata, in a Brazilian savanna. We: (i) evaluated the role of elaiosome in seed removal by P. naegelii; (ii) investigated the fate and viability of removed seeds; (iii) tested if soils associated with P. naegelii nests are nutrient-enriched; (iv) compared seedling survival; and (v) the density of seedling and adult M. serrulata nearby to P. naegelii nests compared with those away from these nests (i.e. controls). 3. Rates of removal of M. serrulata seeds were two-fold higher with elaiosomes than without. The ant attractant oleic acid was the dominant fatty acid in elaiosomes, but it was absent from seeds. Removed seeds are taken into nests, and Tetrazolium tests indicated that 95% of seeds remain viable. Soils associated with P. naegelii nests were not nutrient-enriched, and seedling survival was similar nearby to P. naegelii nests compared with control areas. However, densities of both seedling and adult M. serrulata were higher nearby to P. naegelii nests than in control areas. 4. Our findings show that P. naegelii switches its role from seed predator for most plant species to be the dominant seed disperser for M. serrulata, playing a key role in the distribution of adult plants.     

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.     

Composition of harvested seeds and seed selection by the invasive tropical fire ant, <Emphasis Type="Italic">Solenopsis geminata</Emphasis> (Hymenoptera: Formicidae) in Taiwan

Solenopsis geminata (F.) is an invasive ant that is widely distributed in weedy areas and agricultural fields in Taiwan. Previous studies have found that S. geminata harvests the seeds of numerous plants. In the present study, we further investigated the composition of harvested seeds in ant nests and seed selection by workers. The seed caches in S. geminata nests sampled in eight areas in Taiwan suggest that the seeds harvested by workers were diverse and belonged to 52 plant species in 17 plant families. Twenty-three species (44%) belong to the family Gramineae, and most of the seeds weighed from 0.02 to 2.29 mg, which might suggest that these are the main seeds harvested by S. geminata from their habitat. Ten common species with similar seed weights were used to compare the seed preferences of workers from two experimental sites. The results suggest that the seed preference was different between the two experimental sites. The seeds of Casuarina equisetifolia showed the most obvious difference in seed removal speed, which might suggest that S. geminata potentially prefers the encountered seed species in the habitat. The various plant species in the ant nests and seed preference suggest that fire ants easily accept newly encountered plant species. As more than half of the plant species (52%) and the total seed number (63%) belonged to exotic species, the role of S. geminata might be negative because it tends to harvest exotic seeds and has a high opportunity to improve the establishment of exotic seedlings.     

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.     

The role of ants and mammals in dispersal and post-dispersal seed predation of the shrubs Grevillea (Proteaceae)

The role seed predators play in influencing the dynamics of plant populations has been little studied in Australia. The interaction of ant dispersal and seed predation on the soil seedbank in six shrubby species of Grevillea from the Sydney region of southeastern Australia was examined in selective exclusion experiments, seed array trials and placement of single seeds on the ground.Two distinct seed types in Grevillea were examined and different seed dispersal and post-dispersal seed predation patterns were associated with each: (a) seeds lacking an elaiosome were not attractive to ants and annual seed losses of between 82 and 95% were found in vegetation unburnt for greater than 8 years. Native rodents, Rattus fuscipes, and macropods, Wallabia bicolor, were responsible for these seed losses; (b) seeds with an elaiosome were rapidly handled by ants. Two functional types of ants were recognised. Most encounters were by ants that were small (Local) relative to seed size and these ants simply removed the elaiosome in situ or moved seeds only small distances (<20 cm). Some 0–24% of ant/seed encounters were by large (Removalist) ant species that were capable of moving seeds back to nests. In addition, Rattus fuscipes and Wallabia bicolor consumed at least 32–68% of seeds of Grevillea species with an elaiosome.Ants may reduce the overall levels of seed predation where seeds moved by Removalist ant species escape predation and are deposited in safe sites, hence allowing more seeds to reach the persistent soil seedbank. Mammals do not consume all seeds when ants are excluded, allowing for the potential for some seed escape from predation after seeds are discarded by Local ant species.