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.     

Convergent evolution of seed dispersal by ants,and phylogeny and biogeography in flowering plants: A global survey

Seed dispersal is a fundamental life history trait in plants. Although the recent surge of interest in seed dispersal by ants (myrmecochory) has added greatly to knowledge on the ecology of seed dispersal and ant–plant mutualisms, myrmecochory also represents a unique opportunity to examine the links between seed dispersal and evolution in flowering plants. Here we review the taxonomic, phylogenetic and biogeographic distribution of myrmecochory in flowering plants. Myrmecochory is mediated by elaiosomes, i.e., lipid-rich seed appendages that attract ants and serve as rewards for dispersal. We surveyed the literature for evidence of elaiosomes in angiosperm plants to estimate the global prevalence of myrmecochory. We then searched the literature for phylogenetic reconstructions to identify myrmecochorous lineages and to estimate the minimum number of independent evolutionary origins of myrmecochory. We found that myrmecochory is present in at least 11 000 species or 4.5% of all species, in 334 genera or 2.5% of all genera and in 77 families or 17% of all families of angiosperm plants. We identified at least 101, but possibly up to 147, independent origins of myrmecochory. We estimated three or more origins in 13 families and found that at least half the genera are myrmecochorous in 10 families. Most myrmecochorous lineages were Australian, South African or northern temperate (Holarctic). A mapping of families containing myrmecochorous genera on a dated angiosperm supertree showed that myrmecochory has evolved in most of the major angiosperm lineages and that it is more frequent in younger families (crown group age <80 million years) than in older ones. We suggest that the relatively low physiological and energetic costs of producing an elaiosome and the consistent selective benefits of myrmecochory (dispersal, protection from seed predators and fire, safe and nutrient-rich microsites) explain the numerous evolutionary and developmental origins of myrmecochory in angiosperm plants, and we propose that elaiosomes thus provide one of the most dramatic examples of convergent evolution in biology.     

Dispersal mechanisms of the narrow endemic <Emphasis Type="Italic">Polygala vayredae</Emphasis>: dispersal syndromes and spatio-temporal variations in ant dispersal assemblages

This study assesses the dispersal mechanisms of the narrow endemic Polygala vayredae, analysing the functioning of its dispersal syndromes (anemochory and myrmecochory), the spatio-temporal variability of the disperser assemblage, foraging behaviour and dispersal ability, and the role of the elaiosome in ant attraction and seed germination. The dispersion of diaspores begins when either (1) capsules or seeds fall beneath the mother plant (barochory) or (2) the seeds are directly collected in the suspended capsules by ants (myrmecochory). As capsules frequently open and expose/disseminate seeds before leaving the mother plant, the adaptation for anemochory appears to be reduced and rarely functional, possibly with only occasional events of long-distance dispersal (e.g. under extreme weather conditions). P. vayredae is essentially myrmecochorous and a diverse array of ant species are involved in seed manipulation, with the elaiosome playing a major role in ant attraction. From the plant’s perspective for dispersal, the majority of ant species had a positive interaction with the seeds, but negative and potential neutral interactions were also observed. Overall, dispersal distances were limited and were mainly determined by ant body size. The frequency of interactions and the ant assemblage varied significantly both spatially and temporally, and these factors may have an effect on directing or disrupting the selection of plant traits. Low seed predation and similar germination rates of intact seeds compared with seeds without elaiosome indicate that seed predator avoidance and seed germination improvement after ant manipulation are not among the selective advantages of myrmecochory operating at present. Dispersal mechanisms that enhance seed dispersal within the population and only occasionally lead to long-distance dispersal events, along with the rarity and patchiness of suitable habitats, may be the main factors explaining the actual density and narrow distribution of this species.     

Are subordinate ants the best seed dispersers? Linking dominance hierarchies and seed dispersal ability in myrmecochory interactions

True myrmecochory involves the dispersal of elaiosome-bearing seeds by ants. Between the guild of ants that are attracted to these seeds, only a few of them will act as effective dispersers, that is, transporting the seeds to suitable sites (the nests) for germination and plant establishment. Ant communities are known to be highly hierarchical, and subordinate ants quickly deliver resources to their nest rather than consuming it on-site, thereby avoiding encounters with more dominant species. As a result of a series of studies that were carried out during summer in semi-arid Northwest Argentina, we have found that the most important seed disperser of the myrmecochorous plant Jatropha excisa Griseb. (Euphorbiaceae), the ant Pogonomyrmex cunicularius pencosensis Forel, was the most subordinate species during interspecific interactions. The daily timing of release of the J. excisa seeds through ballistic dispersal increased their probability of being removed by the highly thermophilic P. cunicularius pencosensis. Foraging during the warmest hours of the day allowed P. cunicularius pencosensis ants to avoid the risk of interference competition with dominant species, which also behaved as elaiosome predators. As a conclusion, subordinance behaviour appears to be integral to successful myrmecochory, and also the timing of seed release plays a key role in shaping the dynamics of myrmecochorous interactions. Therefore, ant-dispersed plants should not only favour their discovery by subordinate ants, but also should present their seeds at those times of the day when the behaviourally dominant ants are less active.     

SEED DISPERSAL IN VIOLA (VIOLACEAE): ADAPTATIONS AND STRATEGIES

Surveys of Viola dispersal mechanisms result in the distinction of two major adaptive syndromes: one purely myrmecochorous for the exploitation of seed-transporting ants, the other partially myrmecochorous (= diplochorous) for the explosive ejection of seeds followed by ant exploitation. Diplochory is exhibited by the majority of Viola species, but myrmecochory is exhibited only by eleven species with limited Eurasian distribution. Morphological and experimental evidence suggests the hypothesis that Viola is basically myrmecochorous but that different selective pressures, especially seed predation, have produced a clear divergence in dispersal systems. The majority of species, the diplochores, have evolved a system which combines ballistic and ant seed dispersal with predator avoidance. Diplochory itself may be a response to predation pressure. The minority of species are purely myrmecochorous, possibly highly coevolved with specific ant species, thus limiting the distribution of the Viola species concerned.     

Are gastropods, rather than ants, important dispersers of seeds of myrmecochorous forest herbs?

Seed dispersal by ants (myrmecochory) is widespread, and seed adaptations to myrmecochory are common, especially in the form of fatty appendices (elaiosomes). In a recent study, slugs were identified as seed dispersers of myrmecochores in a central European beech forest. Here we used 105 beech forest sites to test whether myrmecochore presence and abundance is related to ant or gastropod abundance and whether experimentally exposed seeds are removed by gastropods. Myrmecochorous plant cover was positively related to gastropod abundance but was negatively related to ant abundance. Gastropods were responsible for most seed removal and elaiosome damage, whereas insects (and rodents) played minor roles. These gastropod effects on seeds were independent of region or forest management. We suggest that terrestrial gastropods can generally act as seed dispersers of myrmecochorous plants and even substitute myrmecochory, especially where ants are absent or uncommon.     

Migration of an annual myrmecochore: a four year experiment with Melampyrum pratense L.

A seed sowing experiment was conducted in a mixed secondary woodland on acidic soils in NE Germany with Melampyrum pratense, an annual ant-dispersed forest herb which lacks a natural population in the study area, but is abundant in similar habitats. Each set of 300 seeds was sown within one square metre at three sites in 1997, and the development of the populations was recorded from 1998 onward. Additionally, seed fall patterns were studied in a natural population by means of adhesive cardboard. All trials resulted in the recruitment of populations, which survived and increased in both individual number and area, up to the year 2001. Thus, local distribution of Melampyrum pratense is dispersal-limited. Total individual number increased from 105 to 3,390, and total population area from 2.07 to 109.04 m². Migration occurred in all directions. Mean migration rate was 0.91 m per year, and the highest migration rate was 6.48 m. No individual was recorded beyond 7.63 m from the centres of the sawn squares after three years, suggesting exclusive short-distance dispersal. As primary dispersal enables only distances of up to 0.25 m, ants are presumed to be the main dispersal vectors. Despite differences in individual number and colonization patterns, migration rates did not differ significantly between the populations, but were significantly higher in 2001 due to an increased population size. Colonization patterns were characterized by a rapid, negative exponential decrease of population density with increasing distance from the sown plot, suggesting a colonization by establishment of more or less isolated outposts of individuals and a subsequent gradual infill of the gaps between. My results resemble myrmecochorous dispersal distances in temperate woodlands, and migration rates and patterns across ecotones from ancient to recent deciduous forests. They may function as a colonization model of Melampyrum pratense after accidental long-distance dispersal.     

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.     

An update of the world survey of myrmecochorous dispersal distances

We update the global assessment of seed dispersal by ants and test the hypothesis that the body size of seed‐dispersing ant species varies with latitude in the same way as dispersal distance. We compiled all published data about seed dispersal distance by myrmecochory through March, 2011. We then broke the data down by vegetation type, geography and taxonomy. We also compiled data on body size (body length) of the seed‐dispersing ant species from the studies consulted. Based on 7889 observations, the mean dispersal distance was 1.99 m, although the curve has a long tail extending to 180 m. Considering the mean dispersal distance by ant species and study as independent data, the mean dispersal distance was 2.24 ± 7.19 m (n = 183). Shorter distances are associated with smaller ant species, while the tail of the dispersal curve is due to larger ant species. The mean dispersal distance of myrmecochorous seeds dispersed by ants decreased with increasing latitude, but there was no significant relationship between the body size of dispersing ant species and latitude (i.e. myrmecochorous seed‐dispersing ant species do not follow Bergmann's rule). In 1998 we made three predictions: 1) the dispersal distances of the Southern Hemisphere will decrease with as more data from mesophyllous vegetation are obtained; 2) assuming that ant nest density is higher at lower latitudes, the differences in distances between hemispheres would probably decrease with more data; and 3) numerical differences between dispersal distances in mesophyllous and sclerophyllous vegetation zones would increase with more data. The results obtained since 1998 support the only the third prediction. The dispersal distances in mesophyllous vegetation zones are shorter than in the sclerophyllous vegetation zones, and the difference between 1998 have increased. The differences in dispersal distances between hemispheres are consistent with the avoidance of parent–offspring competition (escape hypothesis).     

Ant-mediated seed dispersal contributes to the local spatial pattern and genetic structure of Globba lancangensis (Zingiberaceae)

Globba lancangensis is a typical myrmecochorous perennial herb found in southwest China. In 2002 and 2003, seed dispersal by ants was examined. Twelve ant species were recorded and found to move seeds 0.01-3.35 m. This removal promoted seed dispersal. In 2003, ant exclusion was made in 4 of 8 study plots for 1 year to practically detect the effects of ant dispersal. As a result, nearest neighbor analyses indicated that although ant dispersal could not significantly change the mean nearest neighbor distance, it obviously reduced the clustering degree of seedlings. Spatial genetic structure (SGS) of mature plants in 2003 and sequent seedlings in 2004 were investigated with intersimple sequence repeats. The whole correlagram of all individuals from 2003 showed significant positive autocorrelations between genetic and geographical distances within 4 m, suggesting a patchy structure at such short distances in the studied population. This pattern is likely associated with limited seed dispersal. The comparative analysis of fine-scale correlograms (<4 m) for ant-excluded treatment indicated lacking ants as the primary dispersers could have a similar pattern to ant dispersal presence, which is likely due to the restriction of ant dispersal distance. In conclusion, ant-mediated dispersal contributes to reduce seedling clustering degree and plays a minor role in developing and maintaining the local SGS in G. lancangensis.     

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.     

Non-random dispersal by ants: Long-term field data versus model predictions of population spread of a forest herb

Myrmecochory, i.e. dispersal of seeds by ants towards and around their nests, plays an important role in temperate forests. Yet hardly any study has examined plant population spread over several years and the underlying joint contribution of a hierarchy of dispersal modes and plant demography. We used a seed-sowing approach with three replicates to examine colonization patterns of Melampyrum pratense, an annual myrmecochorous herb, in a mixed Scots pine forest in north-eastern Germany. Using a spatially explicit individual-based (SEIB) model population patterns over 4 years were explained by short-distance transport of seeds by small ant species with high nest densities, resulting in random spread. However, plant distributions in the field after another 4 years were clearly deviating from model predictions. Mean annual spread rate increased from 0.9 to 5.1 m per year, with a clear inhomogeneous component. Obviously, after a lag-phase of several years, non-random seed dispersal by large red wood ants (Formica rufa) was determining the species’ spread, thus resulting in stratified dispersal due to interactions with different-sized ant species. Hypotheses on stratified dispersal, on dispersal lag, and on non-random dispersal were verified using an extended SEIB model, by comparison of model outputs with field patterns (individual numbers, population areas, and maximum distances). Dispersal towards red wood ant nests together with seed loss during transport and redistribution around nests were essential features of the model extension. The observed lag-phase in the initiation of non-random, medium-distance transport was probably due to a change of ant behaviour towards a new food source of increasing importance, being a meaningful example for a lag-phase in local plant species invasion. The results demonstrate that field studies should check model predictions wherever possible. Future research will show whether or not the M. pratense–ant system is representative for migration patterns of similar animal dispersal systems after having crossed range edges by long-distance dispersal events.     

Myrmecochorous adaptations ofCorydalis species (Papaveraceae) in southern Japan

Dispersal morphology based on the myrmecochorous adaptations for predator avoidance of sevenCorydalis species including two varieties are investigated in southern Japan. Three types of myrmecochory were distinguished: myrmecochory with autochory (diplochory), the explosive ejection of seeds followed by ant transportation; myrmecochory with vegetative reproduction, seed-transportation by ants and reproduction by tuber; and pure myrmecochory, seed-transporting by ants only. Diplochory occurs in one winter annual plant, which has explosive capsules, a smooth seed surface, a small elaiosome, long pedicels and large bracts. Myrmecochory with vegetative reproduction occurs in two perennials that reproduce by tuber, although they also produce a small number of seeds with a medium-sized elaiosome. the pedicels and bracts are medium in size. Pure myrmecochory occurs in five annuals or biennials that are characterized by a rough seed surface with a large elaiosome, comparatively high seed production, short pedicels and only small bracts. Diplochory is exhibited by only one species distributed widely throughout Japan. Myrmecochory with vegetative reproduction is exhibited by species mainly distributed in cool-temperate northern Japan, while true myrmecochory is exhibited by the majority ofCorydalis species in warm-temperate Japan.     

Comparison of seed removal by ants in vegetation on fertile and infertile soils

Plants which have seeds adapted for dispersal by ants (myrmecochores) are more common in some vegetation types than others. In Australia, myrmecochores are common in sclerophyll shrublands which are associated with infertile soils but not in forests on fertile soils. One hypothesis to explain this difference is that ants are less abundant or less active in seed removal on fertile soils. A test of this hypothesis was carried out on four sites in the Royal National Park, south of Sydney, by measuring the number of myrmecochorous seeds removed by ants in a 24 h period. It was found that the removal rates of seeds were slightly higher on fertile sites although this difference was not significant. The hypothesis that a lower removal rate on fertile soils could account for the lower percentage myrmecochory was therefore rejected. Apart from the gross removal rate of seeds by ants, other differences in ant behaviour might be important. It is possible that a greater proportion of ant species unfertile soils are seed predators or ineffective seed dispersers, and evidence obtained indicates that this is worthy of further research.     

Combining distances of ballistic and myrmecochorous seed dispersal in Adriana quadripartita (Euphorbiaceae)

The separate contributions of different vectors to net seed dispersal curves of diplochorous systems have rarely been characterised. In Australia, myrmecochory is a common seed dispersal syndrome and in the majority of such systems, seeds are initially dispersed ballistically. We measured ballistic and myrmecochorous seed dispersal distances in relation to canopies of Adriana quadripartita (Euphorbiaceae) and used a simulation model to estimate the net dispersal curve. We also compared seed removal rates and ant abundances under, and outside, plant canopies to examine how foraging patterns by ants may affect net dispersal.Overall ant abundance did not show a significant numerical response to seedfall; however, the abundance of the main seed dispersing ant, Rhytidoponera ‘metallica’ did. Despite this, seed removal rates did not differ significantly between canopy and open locations. Rhytidoponera ‘metallica’ account for 93% of observed seed dispersal events. On average, the ants dispersed seeds 1.54 m and in doing so, moved seed a mean radial distance of 0.76 m away from canopy edges. This contribution to net dispersal distance by ants is considerable since ballistic dispersal moved seeds a median distance of 7.5 cm. Our simulation model indicated that the combination of ballistic and ant seed dispersal is expected to result in seeds being transported a median net radial dispersal distance of 1.05 m from the canopy edge.Thus in this system, an important function of diplochory may simply be to move a higher proportion of seeds from under the canopy of parent plants than is possible by ballistic dispersal alone. This ‘dispersal-for-distance’ may result in reduced parent–offspring competition or may increase the probability that seeds reach rare safe sites for germination and recruitment.     

Seed re-dispersal of four myrmecochorous plants by a keystone ant in central China

Seed re-dispersal following initial harvesting by ants may have important implications for the distribution and fate of myrmecochorous seeds. However, the probability of seed re-dispersed by ants and the effect it may have on subsequent survival appear variable, the functional role of diaspore, disperser and seed predator to the fate of discarded seeds remain unclear. To clarify the ecology, we compared the consequences of seed re-dispersal by a keystone seed-dispersing ant (Myrmica ruginodis Nylander) for four sympatric myrmecochorous plants common to the temperate deciduous forests in Qinling Mountains, central China. Plants varied in the probability of re-dispersal and in elaiosome condition. Ants preferred seeds with residual elaiosomes, while rodents only consumed the two larger-seeded species, regardless of the elaiosome presence. The scattered distribution of discarded seeds increased the probability of ant re-harvesting and, to some extent, reduced rodent predation. Thus, difference in the probability of seed re-dispersal and its subsequent effect on seed fate in relation to ants and rodents was attributed primarily to the elaiosome condition, seed size and seed spatial pattern. The results imply that seed re-dispersal could affect the fitness of plants and ultimately influence the plant abundance and distribution pattern. This highlights the necessity to incorporate re-dispersal into myrmecochory to advance our understanding of the benefits of myrmecochory to plants.     

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.     

Explosive seed dispersal in two perennial Mediterranean Euphorbia species (Euphorbiaceae)

The distance of explosive dispersal, its pattern in time, and the relative importance of autochory have been studied in two diplochorous species: Euphorbia boetica and E. nicaeensis. The seeds of E. boetica released by explosive dispersal reached a median distance of 156 cm and a maximum of almost 8 m, while the distances reached by the seeds of E. nicaeensis were lower: a median of 132 cm and a maximum of 5 m. The differences in explosive dispersal distance between species seem to depend on both seed mass and caruncle retention. The seeds of both species present a caruncle, but in E. boetica this is tiny, and in most cases is shed during the explosion of the capsules. The distances reached by the seeds of these species, dispersed just by capsule explosion, were similar to or greater than the distances to which ants disperse seeds in the Mediterranean sclerophyllous vegetation. Diplochorous plants may maximize either the distance of primary dispersal or that of secondary dispersal. Given that the seeds of E. boetica, that lose their caruncles, are not gathered by myrmecochorous ants, the results suggest that E. boetica maximizes its primary dispersal distance, whereas E. nicaeensis favors its secondary dispersal.     

Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland

The effect of three ant species (Lasius flavus, Formica spp., Tetramorium caespitum) on soil seed bank formation was studied in temperate mountain grassland. Seed removal experiments, analysis of soil seed content and seed survival experiments were carried out to evaluate the influence of ground ants on the seed fate. In the seed removal experiment seeds of 16 species, including 5 species with elaiosome-bearing seeds (myrmecochores), were exposed and their removal followed for 39 h. On average, ants removed 63.8% of myrmecochorous seeds and 10.9% of seeds without adaptation to ant dispersal. Analysis of soil seed content revealed that myrmecochores, in spite of expectations that they would accumulate in nests of seed dispersing ants, were most abundant in the soil of control plots. Evidence on seed relocation to the ant nests was obtained from a comparison of mounds of seed dispersing and seed non-dispersing ant species, as more seeds were found in the mounds of Formica spp. and Tetramorium caespitum (seed dispersers) in comparison with the mounds of Lasius favus (non-disperser).The soil seed bank of the compared microhabitats (control plots and mounds of 3 ant species) differed in their species composition, seed abundance and vertical distribution. The most distinct qualitative differences were between seed flora of control plots and mounds of Tetramorium caespitum. Control plots had approximately 30,000 propagules per m², which was double the number of seeds found in the ant mounds. In control plots, abundance and diversity of seeds steeply declined with depth; this trend was not observed in the mounds probably due to bioturbation. In the seed survival experiment, more seeds (2 out of 3 species) survived in control plots, which may also contribute to the higher seed abundance in this microhabitat.This study showed that seed relocation by ants does not contribute significantly to seed bank build-up at this study site. Ants may, however, increase the regeneration success of myrmecochores, mainly by dispersal for distance and placement in a larger spectrum of microsites, in contrast to species not adapted for myrmecochory.     

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.