Geographic variation in seed removal of a myrmecochorous herb: influence of variation in functional guild and species composition of the disperser assemblage through spatial and temporal scales

The ecological influence of changes in the functional guild and species composition of ant assemblages on ant‐dispersal mutualisms is still poorly known. Using a multi‐scale approach over an 800 km range within the Iberian Peninsula, we tested the hypothesis that variation in seed removal rate was related to functional guild rather than to species composition variations of disperser assemblages in the myrmecochore herb Helleborus foetidus. At least two premises must be confirmed to validate this hypothesis: 1) ant assemblages that are dissimilar in species composition but with similar functional guild composition will not differ significantly in seed removal, and 2) assemblages with different functional guild composition will render different seed removal services. We conducted 3328 ant‐visitor censuses on 462 individual plants to identify both the species composition and functional guild variation of the ant‐disperser assemblage, and the plant seed removal rate. Functional guild composition of the ant assemblage was determined by the proportion of visits of ants acting as legitimate dispersers, facultative dispersers or elaiosome predators. Results showed that ant‐seed dispersal success seemed to be more sensitive to species composition changes of the ant assemblage than to functional guild shifts. However, this sensitivity was scale‐dependent. Thus, at the fine, inter‐individual scale, seed removal covaried with the species and functional guild composition of the ant assemblages; at the inter‐populational scale, differences in seed removal tended to be related to the dissimilarity of the assemblage species composition rather than the assemblage functional guild; finally, inter‐regional differences in seed removal were unrelated to dissimilarities of the ant assemblage composition or functional guild. Though differences in seed removal and the relative frequency of the legitimate dispersers tended to be positively correlated, none of the above premises were fully confirmed in this study. Therefore, our results did not support in full the hypothesis that the variation in seed removal was explained by shifts in functional guild composition, rather than shifts in species composition.     

Adaptive advantages of myrmecochory: the predator-avoidance hypothesis tested over a wide geographic range

The predator‐avoidance hypothesis states that once released from the parent plant, myrmecochorous seeds are rapidly taken by ants to their nests, where they are protected from predators. Previous studies conducted to test this hypothesis have frequently neglected two major aspects necessary for its verification: 1) the influence of processes acting after the seed release and 2) the spatial evenness of such processes. Thus, large‐scale variations in the mechanisms acting beyond seed release, and possibly influencing seed escape from predators, remain poorly documented. Here, we present the results of a post‐dispersal seed‐removal experiment on the myrmecochorous herb Helleborus foetidus, aimed at verifing the predator‐avoidance hypothesis by considering two key post‐release aspects of seed fate: seed destination (dispersed or nondispersed) and seed burial (buried or not buried). Experiments were performed in four different regions in the Iberian Peninsula. After three days of exposure of seeds to the main predator (fieldmice Apodemus sylvaticus), ca 30% of the seeds were removed. Seed destination affected the proportion of seeds escaping predation, but the sign, magnitude and statistical significance of the effect varied among the geographical regions. In the southern region (Cazorla), seeds dispersed in ant nests or intermediate destinations suffered scarcely any predation, but seeds under reproductive‐age plants experienced losses ca 50%. Conversely, in the northern region (Caurel), seeds in nests suffered significantly greater losses than seeds under plants or intermediate destinations, suggesting that nests were especially unsafe destinations. Seed burial had a strong impact on seed escape from predators, and its effect was highly consistent among geographical regions. In view of the consistency of its effect at different spatial scales, seed burial was a more general mechanism for predation avoidance than seed relocation to ant nests, which was habitat‐ and/or ant‐species‐dependent. Our results thus only partially support the predator‐avoidance hypothesis for the evolution of myrmecochory.     

Ant-nest soil and seedling growth in a neotropical ant-dispersed herb

Summary A major hypothesis concerning the benefits of myrmecochory, seed dispersal by ants, to plants is that ant nests are nutrient-enriched microsites that are beneficial to seedling growth. We experimentally test this hypothesis for a neotropical myrmecochore, Calathea ovandensis, asking two questions: 1) is soil of nests of a seed-dispersing ant chemically or structurally distinct from surrounding soils, and 2) do seedlings grow better in soil collected from ant nests than in randomly collected soil? We found that although ant-nest soil was significantly enriched in nitrate-nitrogen, magnesium, iron, manganese, cadmium and percent organic matter compared to randomly collected soil, seedling growth was not significantly improved by ant-nest soil.     

Frugivorous birds mediate sex‐biased facilitation in a dioecious nurse plant

Abstract. Facilitation by dispersal occurs if the nurse plant acts as a focus which is actively selected by seed dispersers and enhances the fitness of the facilitated plant. Sex‐biased facilitation may be produced if seed dispersers tend to concentrate the seeds under female, fruit‐bearing plants of dioecious species more often than under conspecific males. Juniperus sabina is a dioecious shrub with a prostrate growth form from Mediterranean high mountains that modifies many microhabitat characteristics related to seedling establishment and survival. Soil water availability, maximum soil temperature in summer, organic matter and total nitrogen content, were different on open ground as compared with beneath J. sabina shrubs, irrespective of its sex. Other studied characteristics such as soil bulk density and soil compaction after rain did not differ between the microhabitats considered. Some species, such as Juniperus communis, Pinus nigra, Helleborus foetidus and Euphorbia nicaeensis, are spatially associated to J. sabina shrubs, strongly suggesting a facilitative role. The anemochorous P. nigra and myrme‐chorous H. foetidus and E. nicaeensis did not associate preferentially to any sex of J. sabina. Only J. communis, an endozoochorous species sharing the same bird dispersers as J. sabina, presented a female‐biased spatial association with the nurse plant. Seed dispersal mediated by birds attracted by the fruit‐rewarding females of J. sabina explains the sex‐biased spatial pattern of Juniperus communis.     

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.     

Geographic variation in the adaptive landscape for seed size at dispersal in the myrmecochorous <Emphasis Type="Italic">Helleborus foetidus</Emphasis>

Literature on seed dispersal mutualisms suggests that plant populations should hardly adapt to their current dispersers. We address the predictions that selection pressures exerted by ants on dispersal-related diaspore traits of the ant-dispersed Helleborus foetidus are highly variable in space, and that geographic (inter-population) variation in these traits is unrelated to selection by current dispersers. To test these predictions we use the concept of the quantitative adaptive landscape for seed size at dispersal. Such landscape depicts the relationship between the population’s mean trait value (mean seed size in the present study) and the population’s mean fitness (mean dispersal probability in the present study). Adaptive landscapes make it possible to assess whether the mean population’s phenotype agrees with one favored by selection. We first analyse, in 12 populations of H. foetidus from southern Spain, the extent of divergence among populations in seed and elaiosome size, and the abundance, composition, and behavior of the ant communities. Seeds from a fixed set of five of these populations were offered to ants in all the study sites to fit the adaptive landscape for seed size. In addition, seeds from the local population were also offered in each site. Our results show that seed size has undergone a larger divergence among populations than elaiosome size. Despite geographic variation in ant assemblages, the adaptive landscapes for seed size at dispersal were remarkably similar among sites: ants create disruptive selection on seed size in 10 out of 12 study sites. As predicted, the basic features of these adaptive landscapes (curvature and location of the minimum) varied geographically in accordance with variation in the size of seed dispersers. Also as predicted, in most populations, the observed mean seed size does not agree with that expected from the adaptive landscapes at dispersal. However, the relevance of dispersers for seed size evolution should not be neglected since the agreement between observed and optimum seed size was stronger where dispersers were more abundant. Thus, against the general view, our results evidence that, in H. foetidus, the observed geographic variation in dispersal-related plant traits is partly linked to selection exerted by current dispersers. Geographic variation in ant assemblages determines both the existence of a selection mosaic and the degree of adjustment of populations to the patterns of selection in the mosaic.     

The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.     

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.     

Pogonomyrmexcunicularius as the keystone disperser of elaiosome‐bearing Jatropha excisa seeds in semi‐arid Argentina

Myrmecochory or seed dispersal by ants is often described as a diffuse mutualism, because many of the ant species that function as partners are considered to be similar in terms of the frequency and consequences of their interactions. In this work, we test this assumption by conducting ant community surveys and seed removal experiments in six study sites located within a semi‐arid region of northwest Argentina. At each site, we characterized the ant assemblage that interacted with the seeds of Jatropha excisa Griseb. (Euphorbiaceae), an ant‐dispersed native shrub. Our results demonstrate that seed removal was dominated by one species, Pogonomyrmex cunicularius Mayr (Hymenoptera: Formicidae: Myrmicinae), which was responsible of 84% of the observed seed removal events. Although several ant species were attracted to the elaiosome‐bearing seeds of J. excisa, seed removal did not depend on ant community composition (species richness and ant activity) but was significantly influenced by the abundance of P. cunicularius. Its physical, behavioral, and ecological attributes are common with other ant species that have been characterized as keystone seed dispersers in other regions of the world. Nest feeding with marked seeds revealed that once P. cunicularius ants consume the elaiosomes, seeds are left inside the nests undamaged and at an appropriate depth for emergence. Our results support the hypothesis that myrmecochory is often an unevenly diffuse mutualism (i.e., one partner species is particularly important) and that at a local scale P. cunicularius is the keystone seed disperser of J. excisa.     

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

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

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.     

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.     

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.     

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

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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.     

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

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

Geographical variation in the potential of mice to constrain an ant-seed dispersal mutualism

Pre‐ and post‐dispersal Helleborus foetidus (Ranunculaceae) seed predation by mice Apodemus sylvaticus as well as post‐dispersal seed removal by ants was studied, during two years, in six plant populations within three geographical regions (Caurel, Cazorla and Mágina) of the Iberian Peninsula. An observational approach revealed strong interregional differences in seed predation by mice during the pre‐dispersal phase, with high and similar rates of predation in Cazorla and Mágina and much lower rates in Caurel. There were also significant inter‐annual variations on pre‐dispersal seed predation by mice, while the existing habitat‐related differences (of lower magnitude) were not consistent across regions. Field experiments based on seed‐offering exclosures, showed that, despite some interregional variation, post‐dispersal seed removal by ants was consistently high through all spatial and temporal scales considered, with most seeds being removed within 48 h. Conversely, post‐dispersal seed predation by mice was highly variable among regions, being very high in Cazorla and minimal or absent in Caurel and Mágina. Interestingly, in Cazorla, in presence of mice, the number of seeds removed was rather independent of the presence/absence of ants, while under mice exclusion, it was determined by the presence/absence of ants. Conversely, in Caurel, the number of seeds removed by each remover agent (ants or mice) was independent of the presence/absence of the other agent. Thus, though uniquely in Cazorla, mice limited the number of seeds available to ants and, therefore, in this region could potentially have interfered on the development of seed traits that enable ants to efficiently harvest them. Our results support the notion that geographical variation over the Iberian Peninsula of mice seed predation may have promoted a mosaic of well‐matching and mismatching situations between H. foetidus diaspore traits and the characteristics of ant communities, which is consistent with some recent theories on the geographical structure of interactions.     

Reduced dispersal at nonexpanding range margins: A matter of disperser identity

The evolution of dispersal at range margins received much attention recently, especially in the context of dynamic range shifts, such as those following climate change. However, much less attention has been devoted to study variation in and selection on dispersal at nonexpanding range margins, where populations are often small and isolated, and empirical test is dearly missing. To fill this gap, we tested whether dispersal of an ant‐dispersed perennial plant (Sternbergia clusiana) is quantitatively and/or qualitatively reduced toward a nonexpanding range margin. We evaluated plant investment in dispersal structures (elaiosome), seed removal rates, and the relative abundance, activity, and behavior of low‐ and high‐quality seed‐dispersing ants in six sites ranging from mesic Mediterranean site to arid site (>600 to <100 mm of annual rainfall, respectively), which marks the southern range margin of the species. In a set of cafeteria and baiting experiments, we found that overall seed removal rates, the contribution of high‐quality dispersers, maximum dispersal distance and dispersal‐conducive ant behavior decreased toward range margins. These findings agree with a lower investment in reward by range margin plant populations, as reflected by lower elaiosome/seed ratio, but not by variation in the reward chemistry. More than variation in traits controlled by the plants, the variation in ant–seed interactions could be attributed to reduced presence and activity of the more efficient seed‐dispersing ants in the marginal populations. Specifically, we found a mismatch between local distribution of potentially effective seed dispersers and that of the plant, even though those dispersers were observed in the study site. Interestingly, although the observed variation in the outcome of ant–seed interactions supported the prediction of reduced dispersal at nonexpanding range margins with small and isolated populations, the underlying mechanism seems to be incidental difference in the seed‐dispersing ant community rather than a plant‐mediated response to selection.     

Seed dispersal of the hemiparasitic plant Thesium chinense by Tetramorium tsushimae and Pristomyrmex punctatus

Heterotrophic plants normally produce a vast number of dust‐like seeds containing minimal energy reserves, which are usually wind dispersed. However, some heterotrophic species have evolved adaptive strategies to use zoochorous seed dispersal. Seed dispersal by ants, known as myrmecochory, is one of the most widespread animal dispersal systems and has been reported in a diverse range of plant taxa. However, the combination of myrmecochory and heterotrophy seems to be very rare. Here I report the discovery of myrmecochory in the hemiparasitic plant Thesium chinense by Tetramorium tsushimae and Pristomyrmex punctatus. Myrmecochory would be an advantageous dispersal system for T. chinense because its fruits are quickly transferred to the ants' nests, which provide a refuge from the seed predator Canthophorus niveimarginatus. Myrmecochory is also potentially beneficial for T. chinense, as the nests of these ants are frequently located close to poaceous plants, which are the preferred hosts.     

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.