Effects of seed passage through slugs on germination

Seed dispersal by invertebrates is mostly external (e.g. by ants). However, internal dispersal (endozoochory) seems to be uncommon due to size limitations. Slugs are generalist herbivores and increasing evidence suggests that they often disperse seeds. Nevertheless, we know very little on the consequences for plant recruitment. Here, we assess the effect of slug seed passage on germination and early seedling establishment of a set of nine common plant species with limited dispersal capabilities (unassisted or ant dispersed). Germination trials were performed under natural conditions (NW of Spain). Size was a clear limitation for seed ingestion: smaller seeds were eaten more readily, whereas only the largest slugs could swallow the largest seeds. All ingested seeds were voided undamaged. Only not-ingested seeds (the biggest) were damaged, with chewing marks in the surface and 0.8% (N = 250) were broken. In some cases, slugs ate the elaiosomes but discarded the seeds. Slug seed passage had an overall positive effect on total germination and sped it up in two species. Elaiosome removal by slugs had also a positive effect in Ulex europaeus seeds (+160%). However, slugs had no effect on early seedling survival and growth of the study species. Our results show that slugs are effective dispersers in terms of seed treatment in the digestive track. However, according to their reduced movement range (in the range of tens of metres; as shown by other studies), they can only provide rather limited dispersal as compared to vertebrate dispersers. This can be nonetheless significant for species with limited dispersal, e.g. unassisted and ant-dispersed plants, for which they act as non-standard dispersers.     

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

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

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 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.     

Invasive earthworms as seed predators of temperate forest plants

Soil seedbanks play a key role in forest plant communities, contributing to regeneration and acting as a refuge from seed predators. This study provides evidence that seeds entering the soil seedbank are vulnerable to granivory by invasive earthworms in temperate forests. Overall, 73 % of seeds of 6 ecologically important forest species were removed from the soil surface over 2 weeks in a Lumbricus terrestris microcosm experiment; 30 % vanished entirely, and presumably were destroyed. The invasive garlic mustard, Alliaria petiolata, was subject to the highest rates of removal. In contrast, results from a field exclosure experiment using 23 species of seed indicate that while seed predation by worms is still detectable, predation by rodents often may mask impacts of earthworms under natural conditions. Worms and rodents preferred different sizes of seeds: while seed predation by rodents was high in mid- to large-seeded species, earthworms tended to prefer smaller seeds. These findings suggest that although rodents are the main driver of seed predation, invasive earthworms may act as an additional ecological filter, and potentially may further influence the species composition of forest plant communities.     

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.     

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

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

Odor of achlorophyllous plants’ seeds drives seed‐dispersing ants

Seed dispersal by ants is an important means of migration for plants. Many myrmecochorous plants have specialized appendages in their seeds called elaiosome, which provides nutritional rewards for ants, and enable effective seed dispersal. However, some nonmyrmecochorous seeds without elaiosomes are also dispersed by ant species, suggesting the additional mechanisms other than elaiosomes for seed dispersal by ants. The seeds of the achlorophyllous and myco‐heterotrophic herbaceous plant Monotropastrum humile are very small without elaiosomes; we investigated whether odor of the seeds could mediate seed dispersal by ants. We performed a bioassay using seeds of M. humile and the ant Nylanderia flavipes to demonstrate ant‐mediated seed dispersal. We also analyzed the volatile odors emitted from M. humile seeds and conducted bioassays using dummy seeds coated with seed volatiles. Although elaiosomes were absent from the M. humile seeds, the ants carried the seeds to their nests. They also carried the dummy seeds coated with the seed volatile mixture to the nest and left some dummy seeds inside the nest and discarded the rest of the dummy seeds outside the nest with a bias toward specific locations, which might be conducive to germination. We concluded that, in M. humile seeds, volatile odor mixtures were sufficient to induce seed‐carrying behavior by the ants even without elaiosomes.     

Invasive Argentine ants (<Emphasis Type="Italic">Linepithema humile</Emphasis>) do not replace native ants as seed dispersers of <Emphasis Type="Italic">Dendromecon rigida</Emphasis> (Papaveraceae) in California,USA

We investigated the indirect effects of Argentine ant (Linepithema humile) invasions on patterns of seed dispersal and predation in the myrmecochorous tree poppy Dendromecon rigida in coastal San Diego County, California. Significantly more seeds were removed from ant-accessible seed stations at sites numerically dominated by a common harvester ant (Pogonomyrmex subnitidus), a native disperser of these seeds and a species sensitive to displacement by L. humile, than from those stations at sites where L. humile was in the majority. Predation of seeds was high, but variable, across sites, suggesting that reduced dispersal could result in increased seed predation in some habitats. Removal of elaiosomes did not affect the frequency with which predators removed seeds, but ants removed significantly more seeds with elaiosomes than without. In behavior trials, only P. subnitidus was able to carry seeds of Dendromecon rigida effectively. L. humile and a small native ant species, Dorymyrmex insanus, while displaying interest in the diaspores, were seldom able to carry whole seeds and, when they did, only carried them a few centimeters. Displacement of native harvester ants by L. humile appears to decrease the dispersal of Dendromecon rigida seeds and may be increasing loss of seeds due to predation.     

Signals Can Trump Rewards in Attracting Seed-Dispersing Ants

Both rewards and signals are important in mutualisms. In myrmecochory, or seed dispersal by ants, the benefits to plants are relatively well studied, but less is known about why ants pick up and move seeds. We examined seed dispersal by the ant Aphaenogaster rudis of four co-occurring species of plants, and tested whether morphology, chemical signaling, or the nutritional quality of fatty seed appendages called elaiosomes influenced dispersal rates. In removal trials, ants quickly collected diaspores (seeds plus elaiosomes) of Asarum canadense, Trillium grandiflorum, and Sanguinaria canadensis, but largely neglected those of T. erectum. This discrepancy was not explained by differences in the bulk cost-benefit ratio, as assessed by the ratio of seed to elaiosome mass. We also provisioned colonies with diaspores from one of these four plant species or no diaspores as a control. Colonies performed best when fed S. canadensis diaspores, worst when fed T. grandiflorum, and intermediately when fed A. canadense, T. erectum, or no diaspores. Thus, the nutritional rewards in elaiosomes affected colony performance, but did not completely predict seed removal. Instead, high levels of oleic acid in T. grandiflorum elaiosomes may explain why ants disperse these diaspores even though they reduce ant colony performance. We show for the first time that different elaiosome-bearing plants provide rewards of different quality to ant colonies, but also that ants appear unable to accurately assess reward quality when encountering seeds. Instead, we suggest that signals can trump rewards as attractants of ants to seeds.     

Seed dispersal in Erythronium dens-canis L. (Liliaceae): variation among habitats in a myrmecochorous plant

Erythronium dens-canis is a geophyte which produces a single flower each season. The fruits produce small seeds with relatively large elaiosomes. We performed experiments to investigate primary and secondary seed dispersal mechanisms of this species in different habitats in the western part of the Cantabrian Range in northwest Spain. Sticky traps were used to measure primary dispersal of seeds up to 0.5 m from mother plants. Seed cafeteria experiments were performed in different habitats to examine the role of ants and rodents in secondary seed transport and seed predation. Our results indicate that: (a) primary seed dispersal is positively skewed (99% of seeds fall within 20 cm of the mother plant) and seed dispersal distances vary significantly among plants; (b) secondary dispersal is exclusively by myrmecochory, although the proportion of seeds removed by ants differs significantly among habitats; (c) ant species composition and abundances vary among habitats; and (d) freshly dropped seeds are more likely to be removed than seeds that have begun to dry out. We conclude that secondary dispersal of seeds is greatly influenced by habitat but not by small-scale microhabitat. This revised version was published online in June 2006 with corrections to the Cover Date.     

秦岭森林鼠类对华山松种子捕食及其扩散的影响

森林鼠类的种子贮藏行为对植物的扩散和自然更新有着非常重要的影响。通过塑料片标记法,2008和2009年的9月—11月分别在秦岭南坡的佛坪国家级自然保护区内调查了森林鼠类对华山松(Pinus armandii)种子的捕食和扩散,结果显示:森林鼠类对华山松种子有着非常大的捕食压力。在2008年,几乎所有的种子(96.4%)在第3天后就被全部取食,而在2009年,也有将近一半的种子(49.6%)在第3天后被取食。但与此同时,鼠类对华山松种子的扩散也起着非常重要的作用。尤其在2009年,第3天时鼠类最高分散贮藏了17.75%的种子,而且直到第19天后仍然有12.25%被贮藏的种子存活下来。华山松种子在两个年份间的扩散历程有很大差异。在2008年,几乎所有的种子都被鼠类取食,贮藏量非常小;而在2009年,种子被贮藏的比例显著的增加。这个结果可能与种子大小年现象有着十分紧密的联系。2008年是华山松种子的小年,产量非常低。鼠类为了满足其日常的能量需求,只能大量的取食有限的种子,而减少其贮藏量。而2009年是华山松种子的大年,产量非常高。鼠类在满足其日常能量需求的同时,还有大量剩余的种子供其贮藏。     

Fern and bryophyte endozoochory by slugs

Endozoochory plays a prominent role for the dispersal of seed plants, and dispersal vectors are well known. However, for taxa such as ferns and bryophytes, endozoochory has only been suggested anecdotally but never tested in controlled experiments. We fed fertile leaflets of three ferns and capsules of four bryophyte species to three slug species. We found that, overall, spores germinated from slug feces in 57.3 % of all 89 fern and in 51.3 % of all 117 bryophyte samples, showing that the spores survived gut passage of slugs. Moreover, the number of samples within which spores successfully germinated did not differ among plant species but varied strongly among slug species. This opens new ecological perspectives suggesting that fern and bryophyte endozoochory by gastropods is a so-far-overlooked mode of dispersal, which might increase local population sizes of these taxa by spore deposition on suitable substrates.     

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

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

Seed dispersal by ants: behaviour-releasing compounds in elaiosomes

Summary In a study of the biochemical basis of seed dispersal by ants, elaiosomes of Acacia myrtifolia and Tetratheca stenocarpa induced seed collection: intact diaspores and elaiosomes were taken rapidly by ants while most seeds remained on the forest floor. Extracts of elaiosomes (non-polar lipids, polar lipids, and aqueous fractions) were differentially collected by ants. Small pieces of pith impregnated with the polar lipid fraction from elaiosomes of either species elicited a removal rate by ants equivalent to that of intact elaiosomes and significantly higher than that of untreated pith. The non-polar lipid fraction, highest in concentration in elaiosomes of both species, elicited removal that did not differ from that of untreated pith. In T. stenocarpa, however, the aqueous fraction also induced removal equivalent to the polar lipid fraction. 1,2-Diglycerides with unsaturated groups are present in the active polar fractions of both species and unsaturated oleate is the major acid group of the glycerides in elaiosomes. Most oleate-containing compounds tested were taken more rapidly by ants than saturated compounds, and oleic acid, associated with corpse-carrying in ants, induced rapid removal. 1,2-Diolein, but not 1,3-diolein, was taken suggesting that the specific configuration of fatty acid moieties influences collection by ants. We hypothesize that a small suite of oleyl-containing compounds in elaiosomes elicit a stereotyped carrying response by a variety of ants. While the nutrient composition of elaiosomes may provide the underlying selective advantage for ants in seed dispersal, specific compounds may manipulate their behaviour and maximize seed dispersal.     

小果野芭蕉种子散布和不同时空尺度上种子被捕食格局

 以分布在云南西双版纳地区的大型先锋草本植物小果野芭蕉(Musa acuminata)为研究材料,研究其种子初次散布过程和不同时空尺度上种子被 捕食格局。小果野芭蕉的成熟果实有75%在夜间被取食和传播,在白天消失的果实则占25%。蝙蝠是其最主要的种子传播者,鸟类在其种子传播 过程中也起到一定的作用。人工摆放种子试验结果显示小果野芭蕉种子的主要转移者是小型啮齿类(鼠类)和蚁类：在开放处理下3 d后转移率为 86%,排除蚁类(鼠类可进入)处理下种子转移率为69%以及排除鼠类(蚂蚁可进入)处理下种子被转移率为56%。季节、地点和生境均显著影响人工 摆放种子被转移强度：雨季显著高于旱季(p<0.001), 野芭蕉生境显著高于与其相连的自然森林和荒地(p<0.001),在人为干扰较少的补蚌自然 保护区显著低于西双版纳热带植物园和新山,而后两者之间并无显著差异(p>0.05)。同时,地点和生境以及季节、地点和生境都有显著的交互 作用。与相邻的森林和荒地相比,野芭蕉群落中种子被鼠类捕食的强度最大且受蚁类二次转移的比例最少,森林和荒地中种子被鼠类捕食的强 度相对较小且蚁类对种子的二次转移比例较高,从而更好地帮助种子逃避鼠类捕食。因此,依赖于食果动物(主要是蝙蝠, 也包括鸟类)的初次 散布是小果野芭蕉种子逃避捕食的关键。     

Effects of ants,ground beetles and the seed-fall patterns on myrmecochory of Erythronium japonicum Decne. (Liliaceae)

Erythronium japonicum (Liliaceae) inhabits deciduous mesic forests of Hokkaido, northern Japan. Myrmecochory of this species was investigated, especially the dispersal frequency, the effect of seed predators and the seed fall pattern. In the quadrat census using marked seeds of E. japonicum, the ant Myrmica kotokui frequently transported the seeds. However, the frequency of seed removal was low and most seeds were dispersed as little as 1 m or less. The spatial distribution of E. japonicum individuals was nearly random and most seedlings were established 5–20 cm away from the fertile plants, indicating that even this small scale of seed dispersal contributes to avoiding crowding of seedlings. Some arthropods, e.g. springtails, spiders and ticks, hindered seed dispersal by devouring elaiosomes and seeds. Although ground beetle species also damaged seeds and elaiosomes, a few of them exhibited seed removal behaviour. E. japonicum dropped their seeds not all at once but bit by bit, taking 3–6 days to drop all seeds. This seed-fall pattern was effective in raising the frequency of seed removal by ants and reducing seed predation by some arthropods.     

How do primates affect seed germination? A meta‐analysis of gut passage effects on neotropical plants

Biotic seed dispersal is a key process maintaining biodiversity in tropical forests where most trees produce vertebrate‐dispersed seeds. Existing meta‐analyses suggest an overall positive effect of vertebrate gut passage on seed germination, but no significant effects for non‐flying mammals. However, previous meta‐analyses combined rodents (seed predators) and primates (seed dispersers) into the non‐flying mammals category, which may confound specific effects of each group on seed germination. However positive effects of monkeys on germination had previously been found in some studies. Here we disentangle the role of Neotropical primates as contributors to seed dispersal in tropical forests by running a meta‐analysis to determine the overall magnitude of gut passage effects on seed germination percentage and mean time to germination. We also compare effect sizes as a function of different feeding guilds, gut complexities, and seed size. Our results show a strong, positive effect of primates on seed germination percentage and on the number of days to first germination. Strictly frugivorous monkeys, the group most threatened by extinction, showed the highest dispersal quality, increasing germination percentage by 75%. Primates that include insects in their diets had no average effect on germination percentage or time. Gut passage had different outcomes on seeds with different sizes; both large and small seeds showed similar increases in germination percentages after gut passage, but only large seeds germinated faster than control seeds after gut passage. Our results show a relevant role for primates in providing high seed dispersal quality and as drivers of forest regeneration. The combined effects of defaunation and forest fragmentation may result in decreased regeneration of trees, which has the potential to affect negatively both forest structure and ecosystem processes. Finally, we provide general guidelines for standardizing research on seed dispersal by primates. Synthesis Consuming fleshy fruits and dispersing seeds is the main ecological service provided by vertebrates to plants. Vertebrate increases seed germination due to treatment given during digestive system passage. Previous meta‐analyses suggest an overall positive effect of vertebrate gut passage on germination, but no insights are available on its variation among different functional groups of mammals. Our analyses indicated that gut passage by Neotropical primates increased seed germination. Strict frugivores, the ones most threatened by extinction, were the most efficient. Our results show a relevant role for primates in providing high seed dispersal quality and as drivers of forest regeneration, which can be meaningful for conservation in a community scale.     

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.     

The evolution of ant-dispersal in a spring-ephemeral Corydalis ambigua (Papaveraceae): timing of seed-fall and effects of ants and ground beetles

Corydalis ambigua is one of major spring-ephemerals exhibiting ant-dispersal in Hokkaido, northern Japan Seed dispersal in this herb species was observed in a field To attract ants, C ambigua has small spherical seeds with large elaiosomes containing lipids In the quadrat census, 284 (11%) of 2560 seeds were moved away from their mother plants and 131 (46%) of them were recovered into ant nests The mean dispersal distance was 48 3 ± SD 34 4 cm Also the spatial distribution of C ambigua seedlings suggested that the seeds were frequently dispersed by ants The direct observations revealed that the main dispersal ant species were Myrmica kotokui and Lastus japomcus Additionally some ground beetles including Plerostichus spp often devoured elaiosomes and seeds Corydalis ambigua seeds were dropped for about two weeks in late May The collecting by pitfall trap revealed that, in this short period, the frequency of ant foraging on the ground was high and ground beetles were a little yet Moreover, in the seed removal experiments, seeds were more frequently removed in this seed-fall period while seed predation by ground beetles was intensive after the period