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.     

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.     

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.     

Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Dispersal and the dynamics of the soil seed-bank

Genet survival in seeds of Acacia suaveolens was examined through both dispersal and dormancy in the soil in populations near Sydney. Following initial passive seed-fall, the majority of seeds lie within a 1 m radius of the stem of the parent. Further dispersal is predominately mediated by ants. A. suaveolens seeds possess an elaiosome which attracts ants. When elaiosomes are removed, the potential for further dispersal of seeds is greatly reduced. Three species of ant disperse seeds of A. suaveolens and the fate of seeds following ant dispersal was observed to depend on the particular species of ant involved. Ants of both Iridomyrmex sp. and Pheidole sp. B are too small to drag seeds and, instead, ants of these species usually remove the elaiosome in situ, with little dispersal of the-seed resulting. Ants of Pheidole sp. A are larger and disperse seeds further, frequently taking them into their nests where the elaisosome is removed. Seeds are retained inside the nests and incorporated into the floors and walls of passageways and chambers. Several supposed ‘advantages’ of myrmecochory were examined but none were verified. Instead, two distinct ‘disadvantages’ were identified. These were: burial of seeds by ants of Pheidole sp. A into ‘unsafe sites’; and too deep a burial of seeds in nests for seeds to receive a stimulus to germinate during fires, and for seedlings to emerge successfully. Outside nests of Pheidole sp. A. seeds are concentrated in the top 5 cm of the soil, whilst within nests of these ants, seeds are found up to 15 cm deep. The dynamics of various components of the soil seed-bank were examined using seeds buried in nylon mesh containers. The seed-bank is persistent without annual recruitment to seedlings, enabling a population to persist as seeds after all above-ground plants have perished.     

A stochastic model of seed dispersal pattern to assess seed predation by ants in annual dry grasslands

A combined field experiment and modelling approach has been used to provide evidence that ants may be responsible for an observed lower patchiness and higher plant diversity in the neighbourhood of ant nests, within Mediterranean dry grasslands belonging to the phytosociological class Tuberarietea guttatae. The hypothesis was that seeds occurring in clumps may have a higher probability to be harvested than seeds having a scattered distribution. In order to test this hypothesis, four analysis steps were performed. First, pattern of seed production and dispersal of four species was recorded; two of them were more abundant next to ant nests (Tuberaria guttata, Euphorbia exigua), whereas the other two were more abundant away from ant nests (Bromus scoparius and Plantago bellardi). Second, a stochastic model was developed to simulate the observed dispersal patterns of each studied species. Third, 10 seed spatial arrangements in accordance to the distribution patterns created by the model were offered to ants and the location of predated seeds was recorded. Finally, the observed pattern of seed predation was matched to models performed by different distributions of probability. Results showed that the probability of being predated decreased as distance among seeds increased. This preference of ants for high concentration of food items holds down the dominant species sufficiently to allow the subordinates to survive, thus increasing diversity near nests. The observed higher frequency of small-seeded, small-sized, or creeping therophytes close to the ant nests can be therefore seen as an example of indirect myrmecophily.     

ANT DISPERSAL OF CALATHEA (MARANTACEAE) SEEDS BY CARNIVOROUS PONERINES (FORMICIDAE) IN A TROPICAL RAIN FOREST

Two tropical herbaceous plant species, Calathea microcephala and Calathea ovandensis, exhibit morphological adaptations for seed dispersal by ants. In the field, 21 ant species are attracted to the arils of the seeds. Previously known as predatory carnivores, the ponerine ants Odontomachus laticeps, O. minutus, Pachycondyla harpax, and P. apicalis carry the seeds toward their nests, behaving as though they were carrying prey. These ants remove the aril in the nest, and seeds without arils germinate more readily than seeds with arils. Ant foraging distances can account for seedling distributions. Myrmecochory may be far more common in tropical rain forests than has been previously suspected.     

Seed harvesting by ants in forests of Eucalyptus regnans F. Muell. in central Victoria

In an unburnt, mature forest the germination of E. regnans on undisturbed bare ground is very poor in spite of adequate seed fall, and the presence of conditions suitable for germination over much of the year. No seed storage occurs in the top soil in spite of some temporary seed dormancy and the disturbance of the surface soil by earthworms and lyre birds. Seed is removed from the top soil by several ant species– Prolasius pallidus, P. brunneus, P. flavicornis and Chelaner leae –and is taken into nests. Removal of seed is more rapid and complete in summer than winter and is more vigorous in young forest than old. Observations on artificial nests in the field and laboratory indicate that seed is eaten and not stored for any length of time. The testae may be left intact, fragmented or moulded into crumbs with other material. Certain sugar-like substances, which are extractable in low concentrations from fresh seed, may be involved in the attractivity of seed to ants. The numbers of ant species in the forests vary with the micro-climate of the site, and in the mature understorey of Pomaderris aspera the activities of the three dominant species Iridomyrmex biconvexus, Prolasius pallidus and P, hrunneus are largely separated diurnally, seasonally, and spatially in the foraging areas of the litter layer. The density of the nests of seed-harvesting ants is high, particularly in bare areas of the forest floor. The total number of ants probably exceed 5–6 million/ha and is probably sufficient to effectively remove 60% of the sporadic seed fall of normal years. It is suggested that the success of germination of E. regnans after wild fires is not due to any specific stimulation but rather to a temporary interference of ant foraging activity and then to the saturation of their food requirements by a massive seed fall from canopy-stored capsules.     

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.     

Harvester ant foraging and plant species distribution in annual grassland

Summary The harvester antVeromessor andrei Mayr is a major seed predator on annual grassland growing on serpentine soil at Jasper Ridge, N. California. Ants forage intensively during morning and evening sessions in areas surrounding nests. Activity is at its most intense in early summer, coinciding with peak seed release for most annual plant species. Ants show strong preferences for seeds of non-dominant species, notablyMicroseris douglasii (DC.) Sch.-Bip., but preferences alter over the season in response to seed availability. Seeds of the dominant annual species,Lasthenia californica DC ex Lindley are not foraged until later in the summer when seeds of other species are less abundant.Seedling densities and species compositions on ant nests differ markedly from surrounding areas with species relative abundances being similar to those found on gopher mounds. An exclosure experiment in areas adjacent to nests indicated that ants significantly reduced the densities of species with preferred seeds. Ants may therefore significantly affect plant distribution and abundance within the serpentine grassland.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Interaction between ants and fruits of<Emphasis Type="Italic"> Guapira opposita</Emphasis> (Nyctaginaceae) in a Brazilian sandy plain rainforest: ant effects on seeds and seedlings

This study examines the dispersal system of Guapira opposita in a tropical sandy rainforest in southeast Brazil. Guapira trees produce small fruits with a high protein content (28.4%) and low lipid content (0.3%), and the plant is primarily dispersed by birds. Mature fruits of G. opposita can fall spontaneously with the pulp intact, or be dropped by birds with bits of pulp attached. In either case, ground-dwelling ants rapidly remove the fruits to their nest (93% after 12 h). The ponerine ants Odontomachus chelifer and Pachycondyla striata are the main seed vectors among the ants, and together account for 56% (20 of 36) of the ant-fruit interactions recorded on the forest floor. Individual workers of O. chelifer and P. striata transport single fruits to their nests. Bits of pulp are fed to larvae and worker nestmates, and intact seeds are discarded outside the nest. Germination success in Guapira is higher for cleaned seeds (pulp removed) than for seeds coated by pulp. Guapira seedlings and juveniles are more frequent close to Odontomachus nests than at sites without such nests. Soil samples from Odontomachus nests had greater penetrability, and higher concentrations of P, K, and Ca than random soil samples. Field experiments suggest that the association between G. opposita seedlings and O. chelifer nests can potentially render the plant some protection against herbivores. Results indicate that fruit displacement by ponerine ants play an important role in the biology of G. opposita seeds and seedlings in the sandy forest, and illustrate the complex nature of the dispersal ecology of tropical tree species.     

Impacts of Argentine ants on avian nesting success

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

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

相似文献   

Myrmecochory in Polygala vulgaris L., Luzula campestris (L.) DC. and Viola curtisii Forster in a Dutch dune area

Summary Seed dispersal by ants in Polygala vulgaris, Luzula campestris and Viola curtisii was studied in a primary dune valley on the island of Terschelling, The Netherlands. Normally developed seeds of all three species are taken by the ants into their nests. The ants show a distinct preference for the seeds of the specialized myrmecochore Polygala vulgaris, as compared with the two diplochorous species. It could be demonstrated that the elaiosome is the attractive part of the seed. Mapping studies demonstrate that the dispersal of the seeds by ants has a marked effect on the distribution pattern of the standing population of Polygala and Viola. Adult plants are often found on or close to the active nest mounds of all ant species present, while the growing sites of juvenile individuals and seedlings are practically restricted to the nest environment. The nests of two of the seed-dispersing ant species, viz., those of Lasius niger and Tetramorium caespitum, show differences in soil chemistry with the surroundings. The ant nests are significantly richer in some essential plant macronutrients, such as phosphate, potassium and nitrate. The advantage of myrmecochory in the dune area of Terschelling is 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.     

Arboreal ants as key predators in tropical lowland rainforest trees

Ants numerically dominate the canopy fauna of tropical lowland rain forests. They are considered to be key predators but their effects in this regard have only rarely been studied on non-myrmecophytes. A conspicuously low abundance of less mobile, mainly holometabolous arthropods like Lepidoptera larvae corresponds with ant dominance, while hemimetabolous highly mobile nymphs occur regularly and in large numbers in the trees. This is in contrast to the temperate regions where ants are mostly lacking on trees and holometabolous larvae are frequent. In this study we experimentally measured ant predation in the trees by offering caterpillars as baits. Fifty-four ant species were tested, of which 46 killed caterpillars and carried them away to their nests while only eight species ignored the offered larvae. Insecticidal knockdown fogging of ten trees after finishing the prey experiments showed that on average 85% of ant individuals per tree were predacious. With the analysis of another 69 foggings and meticulous observations in many other trees this suggests that arboreal ants are responsible for the low abundance of less mobile arthropods in tropical lowland rain forest canopies. Ant predation was significantly lower in a disturbed forest indicating that human disturbance induces a change in the functional interactions in these ecosystems.     

Predation risk alters interactions among species: competition and facilitation between ants and nesting birds in a boreal forest

Although interactions between species are often assumed to be fixed, theory and empirical evidence suggest that they may be quite variable, changing in the presence of other species or environmental conditions. The interaction between ants and nesting birds exhibits such variability, ants sometimes being predators of bird nests and other times protectors of them. Hypothesizing that predation risk might be a critical factor in altering the interaction of ants with birds, I investigated the interaction of wood ants Formica aquilonia with nesting birds under different levels of predation risk. In a controlled field experiment, I allowed tits ( Parus major, P. caeruleus, P. ater ) and pied flycatchers ( Ficedula hypoleuca ) to select nest boxes in trees with ants (ant trees) or trees without ants. I found that birds usually nested in trees without ants, apparently to avoid the danger of injury from encounters with ants. Nesting in ant trees occurred mainly in the habitat where risk of predation was highest (along the forest edge), and with the bird taxa that lost nests most frequently in trees without ants (tits). Tits nesting on the forest edge achieved significantly greater nesting success, and fledged significantly more young, in ant trees compared with trees without ants. As the season progressed, ant traffic increased in trees without nesting birds, but decreased in trees with nesting birds, indicating that the outcome of interference competition between ants and nesting birds was reversed under increased predation risk. These results support the idea that predation risk can shift species interactions from predominately competitive processes to predominately facilitative processes.     

Invasional 'meltdown' on an oceanic island

Islands can serve as model systems for understanding how biological invasions affect community structure and ecosystem function. Here we show invasion by the alien crazy ant Anoplolepis gracilipes causes a rapid, catastrophic shift in the rain forest ecosystem of a tropical oceanic island, affecting at least three trophic levels. In invaded areas, crazy ants extirpate the red land crab, the dominant endemic consumer on the forest floor. In doing so, crazy ants indirectly release seedling recruitment, enhance species richness of seedlings, and slow litter breakdown. In the forest canopy, new associations between this invasive ant and honeydew‐secreting scale insects accelerate and diversify impacts. Sustained high densities of foraging ants on canopy trees result in high population densities of host‐generalist scale insects and growth of sooty moulds, leading to canopy dieback and even deaths of canopy trees. The indirect fallout from the displacement of a native ‘keystone’ species by an ant invader, itself abetted by introduced/cryptogenic mutualists, produces synergism in impacts to precipitate invasional ‘meltdown’ in this system.     

Comparison of insects and vertebrates as removers of seed and fruit in a Western Australian forest

Removal from the forest floor of seed or fruit of the six commonest species of trees in upland areas of the northern jarrah forest of Western Australia was studied experimentally. The smallest seeds (Eucalyptus marginata, Allocasuarina fraseriana) were taken by insects (presumably ants). Heavier seeds or fruits (E. calophylla, Banksia grandis, Persoonia longifolia, P. elliptica) were taken more often by vertebrates (presumably srrmll mammals). Seeds of E. calophylla and B. grandis were most preferred by vertebrates, possibly because they have the highest relative concentrations of nitrogen and phosphorus.     

Analysis of how ant behaviors affect germination in a tropical myrmecochore Calathea microcephala (P. & E.) Koernicke (Marantaceae): Microsite selection and aril removal by neotropical ants,Odontomachus, Pachycondyla,and Solenopsis (Formicidae)

Summary The evolutionary effects of a tropical ant-seed interaction are examined by posing questions about the fate of Calathea seeds carried by neotropical ants. Where do ants take seeds and what do they do with them? How do ant behaviors affect seed germination? Treatment of seeds by ants is determined by a series of seed-fate trials in captive colonies. There is no evidence of seed predation by ants. Odontomachus laticeps, Pachycondyla spp, and Solenopsis geminata rapidly displace seeds to ant nests, determine the microsites of seeds, and remove the seed arils for food. The seed arils are rich in lipids. The effects on germination of microsite selection and aril removal are quantitatively evaluated. Seeds which are immediately taken to a consistently moist spot germinate readily; 72% germinate, with a mean germination speed of 29 days. For such seeds aril removal does not significantly affect germination. In contrast, seeds which experience a delay before encountering appropriate germination conditions seem to exhibit an induced dormancy (sensu, Harper 1977) and a lower germination percentage. They take longer to germinate (up to 85 days) even after conditions become appropriate. It appears that their germination is enhanced by aril removal, which may act as an environmental cue to break dormancy. Such a mechanism would indicate that ant-handling of seeds is predictive of favorable conditions for seedling growth and establishment. The exact nature of such conditions and the effects on plant population dynamics remain to be seen.