Competitive and allelopathic effects of garlic mustard (<Emphasis Type="Italic">Alliaria petiolata</Emphasis>) on American ginseng (<Emphasis Type="Italic">Panax quinquefolius</Emphasis>)

The purpose of this study was to test competitive and allelopathic effects of invasive garlic mustard on American ginseng (Panax quinquefolius) seedlings under natural conditions. For comparative purposes, we also examined the potential effects of the native striped violet (Viola striata). In order to partition effects of resource competition and chemical suppression via allelopathy, field soils were amended with activated carbon or left unamended. Activated carbon positively affected ginseng growth as well as biomass of competitors. Ginseng mortality tended to increase with garlic mustard presence, though activated carbon alleviated this response. Garlic mustard had no significant effect on ginseng seedling growth, while striped violet suppressed shoot length in the absence of activated carbon. Our results showed a surprising effect of activated carbon on plant growth, a potential allelopathic effect of the native striped violet and suggest that newly invaded ginseng populations with low densities of garlic mustard may be able to withstand its effects. However, recruitment within invaded populations may decline.     

Patterns of seed mass variation and their effects on seedling traits inAlliaria petiolata (Brassicaceae)

Seed mass is considered to be the least plastic component of reproductive yield. Yet, in invasive populations of garlic mustard, Alliaria petiolata, seed mass was highly variable (eightfold among populations, 2.5-7.5 fold within populations, two-threefold within individuals, and 1.4-1.8 fold within fruits). Variation in seed mass among populations explained nearly half of the total variance. Variation among seeds within fruits accounted for a further 25% of variance. Individual seed mass within a plant decreased with increased distance from the main stem, suggesting that access to parental resources limits seed size in a predictable manner. MANOVAs and Roy-Bargmann stepdown analyses revealed significant effects of seed mass, but not seed position (within a fruit or within an infructescence), on an array of subsequent seedling traits. Smaller seeds germinated significantly earlier, and seedlings from small seeds produced their first primary leaves significantly later and grew significantly taller. After accounting for seed mass as a covariate, only one seedling trait, date of first leaf emergence, was affected by seed position in a fruit. Differences in seed mass may therefore affect seedling recruitment via effects on early seedling growth in this weedy species.     

Infection by powdery mildew Erysiphe cruciferarum (Erysiphaceae) strongly affects growth and fitness of Alliaria petiolata (Brassicaceae)

Garlic mustard (Alliaria petiolata) is an invasive biennial that negatively impacts plant and animal communities throughout North America and lacks significant herbivory in its invasive range. Throughout Ohio, many garlic mustard populations support the powdery mildew fungus Erysiphe cruciferarum, although disease incidence varies among populations and environments. Effects of infection on plant growth, as well as both plant and fungal responses to drought and light conditions, were examined on greenhouse-grown, first-year garlic mustard plants. Also, the effects of the fungus on plant growth and fitness were studied in a naturally growing population of second-year plants in the field. Powdery mildew significantly reduced growth of first-year plants in the greenhouse, eventually causing complete mortality. Simulated drought slowed both plant growth and disease development, independent of light conditions. In the field, plants with little incidence of disease after their first year grew taller during their second year, producing significantly more siliques and twice as many seeds as heavily diseased plants did. Seed germination rates did not differ between plants with different levels of disease severity. Consistent reductions in survival, growth, and fitness caused by fungal infection may reduce populations of garlic mustard. These effects may be more evident in moist sites that favor fungal development.     

Comparative seed germination ecology of Austrostipa compressa and Ehrharta calycina (Poaceae) in a Western Australian Banksia woodland

Austrostipa compressa, a native ephemeral of southwest Western Australia was stimulated to germinate under a range of temperatures, in the presence of light, and exposure to smoke-water. This combination of environmental cues results in winter-maximum germination in immediate postfire and disturbed-soil environments of this Mediterranean-type climate. In contrast, Ehrharta calycina, an introduced perennial grass from southern Africa that has invaded Banksia woodlands, germinated under a wide range of temperature and light conditions, but showed no promotive response to smoke-water. Although A. compressa seeds tolerated heat shock better than E. calycina, the self-burial mechanism of A. compressa seeds ensures protection from fire. High-intensity fire could have a greater impact on E. calycina, as the seeds of this species tend to accumulate in the top of the soil profile where they are more susceptible to high temperatures. Although seeds of E. calycina are more susceptible to high temperatures, survival of mature individuals by postfire resprouting ensures continued survival in native woodlands. Estimates of soil seed bank densities showed extreme variability, but some recently burnt areas of the Yule Brook Botany Reserve contained up to 8000 seed m^?2 of A. compressa and nearly 75 000 seeds m^?2 of E. calycina. Viable soil seed bank densities of A. compressa are reduced with time-since-last fire, but areas of greater than 45 years since the last fire, still contained up to 119 seeds m^?2. In both species, only about half their soil seed bank germinates following fire, thus ensuring the potential for later recruitment. Massive soil seed populations of E. calycina in native Banksia woodlands pose a major problem to management of this plant community type.     

Relative contribution of lipid sources to eggs of lesser scaup

Studies of how birds mobilize nutrients to eggs have traditionally considered a continuum of possible allocation strategies ranging from income breeding (rely on food sources found on the breeding grounds) to capital breeding (rely on body reserves stored prior to the breeding season). For capital breeding, stored body reserves can be acquired either on or away from the breeding grounds, but it has been difficult to quantify the relative contribution of each, precluding identification of key habitats for acquiring nutrients for clutch formation. During 2006–2009, we explored the importance of spring‐staging habitats versus breeding‐area habitats for egg‐lipid formation in female lesser scaup Aythya affinis using stable carbon (δ¹³C) isotope analyses. Although δ¹³C values for abdominal lipid reserves brought to the breeding grounds overlapped those of local amphipods, we were able to quantify the importance of local plant carbohydrates (seeds of emergent wetland plants) to the production of eggs. We compared the importance of local wetland seeds (overall δ¹³C: ?29.1 ± 0.9‰ SD) to combined lipid stores and lipids from local amphipods (overall δ¹³C: ?23.8 ± 2.2‰). Local seeds and stored body lipids contributed equally to egg lipid formation across years but we found evidence of annual variation in their relative importance. Wetland seeds contributed 39% (SE = 10%) to egg lipid production, and the importance of this source varied by year (90% CI = 47–75% in 2006, 13–42% in 2007, 29–65% in 2008, and 7–30% in 2009). In contrast to earlier studies that suggest lesser scaup predominantly employ a capital breeding strategy, our results suggest that in some years females may attain half of their energy for clutch formation from foods on the breeding grounds.     

Low allelochemical concentrations detected in garlic mustard-invaded forest soils inhibit fungal growth and AMF spore germination

Garlic mustard’s (Alliaria petiolata, Brassicaceae) invasive success is attributed in part to its release of allyl isothiocyanate (AITC) into the soil. AITC can disrupt beneficial arbuscular mycorrhizal fungi (AMF) associated with native plant roots, which limits their soil resource uptake. However, AITC and its precursor, sinigrin, have never been detected in garlic mustard-invaded forest soils. Here, we use high performance liquid chromatography (HPLC) and gas chromatography-mass spectrometry (GC–MS) to assess the concentration and bioactivity of these putative allelochemicals in paired forest plots uninvaded or invaded by garlic mustard. Our methods detected AITC and sinigrin only where garlic mustard was present and our recovery of AITC/sinigrin coincided with adult senescence. A bioassay of in situ fungal hyphae abundance revealed significantly reduced hyphal abundance in the presence of garlic mustard relative to uninvaded soils. Finally, the lowest concentration of AITC measured in the field (~0.001 mM) is highly inhibitory to the spore germination of a forest AMF species, Glomus clarum. Together, our data provide the first direct evidence of garlic mustard-produced sinigrin and AITC in forest soils and demonstrate that even low levels of these chemicals have the potential to significantly suppress AMF growth and spore germination, strengthening their status as allelopathic novel weapons.     

Buried seed population in the herbaceous lomas on Loma Ancon in the coastal desert of central Peru

Herbaceous lomas in the Peruvian coastal desert, of South America establish in spring, and its habitat is limited to the southern or southwestern slopes along the coast that are affected by thick fog. The time of appearance, the duration and the thickness of the fog vary greatly from year to year, so the lomas can grow only in habitats with enough water to, sustain seed germination and plant growth. This paper studies the species composition and density of the buried seed population, of the herbaceous lomas of Loma Ancon in order to clarify the mechanisms of the lomas' establishment. The mean number of species with viable seeds was about, 12 spp. m⁻² and that of dead seeds was about 22 spp. m⁻². The dominant species wereSolanum tuberiferum, S. pinnatifidum andNolana humifusa, both in viable and dead seeds. Viable seed density was about 5000–8000 seeds m⁻², which is comparable with the seed densities of other herbaceous communities. Dead seed density was about 15000–27000 seeds m⁻², or nearly three times the viable seed density, because the rate of decomposition was slow in the extremely dry conditions. The net increase of viable seeds by seed production was estimated at about 5000 seeds m⁻² in 1980, and the increase in the number of dead seeds was 2200 seeds m⁻².     

The impact of global warming on germination and seedling emergence in Alliaria petiolata,a woodland species with dormancy loss dependent on low temperature

• The impact of global warming on seed dormancy loss and germination was investigated in Alliaria petiolata (garlic mustard), a common woodland/hedgerow plant in Eurasia, considered invasive in North America. Increased temperature may have serious implications, since seeds of this species germinate and emerge at low temperatures early in spring to establish and grow before canopy development of competing species.
• Dormancy was evaluated in seeds buried in field soils. Seedling emergence was also investigated in the field, and in a thermogradient tunnel under global warming scenarios representing predicted UK air temperatures through to 2080.
• Dormancy was simple, and its relief required the accumulation of low temperature chilling time. Under a global warming scenario, dormancy relief and seedling emergence declined and seed mortality increased as soil temperature increased along a thermal gradient. Seedling emergence advanced with soil temperature, peaking 8 days earlier under 2080 conditions.
• The results indicate that as mean temperature increases due to global warming, the chilling requirement for dormancy relief may not be fully satisfied, but seedling emergence will continue from low dormancy seeds in the population. Adaptation resulting from selection of this low dormancy proportion is likely to reduce the overall population chilling requirement. Seedling emergence is also likely to keep pace with the advancement of biological spring, enabling A. petiolata to maintain its strategy of establishment before the woodland canopy closes. However, this potential for adaptation may be countered by increased seed mortality in the seed bank as soils warm.

     

Seed dispersal potential by wild mallard duck as estimated from digestive tract analysis

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Composition of soil seed banks in southern California coastal sage scrub and adjacent exotic grassland

Soil seed banks are important to many plant communities and are recognized as an important component of management plans. Understanding seed bank composition and density is especially important when communities have been invaded by exotic species and must be managed to promote desirable species. We examined germinable soil seed banks in southern California coastal sage scrub (CSS) that is heavily invaded by exotic grasses and in adjacent exotic grassland. Soils from both communities had similar seed banks, dominated by high densities of exotic grass and forb species. Up to 4,000 exotic grass seeds and at least 400 exotic forb seeds/m² were found in most soils, regardless of aboveground vegetation type. Native forbs averaged 400 seeds/m² in grass-dominated areas and about 800 in shrub-dominated soils. Shrub seed density was <1 and <10 seeds/m² in grass- and shrub-dominated areas, respectively, indicating that the shrub seed bank is not persistent compared to annuals. We also compared pre- and post-burn soil seed banks from one location that burned in October 2003. Late-season burning in both grass- and CSS-dominated areas disproportionately reduced exotic grass seed densities relative to native seed densities. The similarity of the seed banks in adjacent grass and shrub communities suggests that without intervention, areas currently dominated by CSS may become more similar to grass-dominated areas in terms of aboveground vegetation. In such areas, the first growing season following a wildfire is a window of opportunity for increasing native diversity at a time when density of exotic grass seeds is low. At time of research, Robert D. Cox was graduate student.     

Allelopathic inhibition of germination by Alliaria petiolata (Brassicaceae)

Garlic mustard (Alliaria petiolata, Brassicaceae) is an invasive, nonindigenous species currently invading the understory of North American woodlands where it is a serious threat to the native flora. Part of this success might be due to allelopathic interference by garlic mustard. Two congeneric species, the European Geum urbanum and the North American Geum laciniatum, were tested for allelopathic inhibition of germination by garlic mustard. Seeds were germinated either on substrate contaminated by garlic mustard or on substrate with contamination neutralized by activated carbon. Allelopathic effects of native European and invasive North American garlic mustard populations were also compared. Activated carbon increased germination by 14%, indicating that garlic mustard contaminated the substrate through root exudates. Activated carbon in turn counteracted this effect. The two test species differed in their sensitivity to allelopathic interference. North American G. laciniatum had a much stronger increase in germination when activated carbon was added to the substrate, independent of the origin of garlic mustard. In contrast, the European G. urbanum germinated better in substrate precultivated with North American garlic mustard, whereas activated carbon increased its germination only in substrate precultivated with European garlic mustard.     

Leaf-cutting ant nests support less dense and impoverished seed assemblages in a human-modified Caatinga dry forest

Regenerating forests make up an increasingly large portion of tropical landscapes worldwide and regeneration dynamics may be influenced by leaf-cutting ants (LCA), which proliferate in disturbed areas and collect seeds for fungus culturing. Here, we investigate how LCA influences seed fate in human-modified areas of Caatinga dry forest. We evaluate the seed deposition and predation on Atta opaciceps nests, foraging habitat surrounding nest and control habitat away of nest influence of 15 colonies located along a forest cover gradient during the rainy and dry seasons. For each habitat, four 50-cm² plots were established and all seeds on the soil surface were collected along 1 year. We recorded 13,628 seeds distributed among 47 species and 36.57% of the total seeds did not show any sign of predation. Nest mound habitats supported low-density and species-poor seed assemblages, which were taxonomically distinct from the control habitats. These effects only occurred in the rainy season. The proportion of undamaged seeds were similar across the habitats. While forest cover did not influence seed assemblage in terms of species richness or seed predation, it did interact with habitat type via increments in seed abundance as forest cover increased across the nests. Forest cover also affected seed composition, but only in the rainy season. These results indicate that LCA decrease seed deposition in areas under their influence, particularly on the nest mounds. As LCA profit from human disturbance in the Caatinga, their role as seed ‘sinks’ should be enhanced in disturbed Caatinga patches, particularly during the rainy season, when most of the plant recruitment occurs. Our findings reinforce the importance of LCA as drivers of forest dynamics and resilience in human-modified landscapes.     

Competitive Interactions among First‐Year and Second‐Year Plants of the Invasive,Biennial Garlic Mustard (Alliaria petiolata) and Native Ground Layer Vegetation

We studied the effects of hand weeding of second‐year plants of the biennial garlic mustard (Alliaria petiolata) on first‐year plants (seedlings) and native ground layer vegetation. Garlic mustard is a Eurasian species that has invaded deciduous forest ground layers in eastern North America. Treatments consisted of a control and an early or late weeding of second‐year garlic mustard. The early treatment (early March) was applied before garlic mustard seeds had germinated and when most native species were dormant. The late treatment (mid‐May) occurred after plants had bolted, flowering was occurring, and most native species and new garlic mustard seedlings were actively growing. Pre‐treatment data were obtained in 2004 and treated and control plots were sampled in 2005, 2006, and 2007. No significant treatment effects were observed in 2004 or 2005. In 2006, mean cover of first‐year plants was higher in the early weeding treatment than in the late weeding treatment and control. In 2007, mean cover of first‐year garlic mustard was higher in the control than in either of the two weeding treatments. There were no significant treatment effects in any year on native vegetation cover, bare ground, or the five most abundant native species. Our data indicate that (1) late weeding of garlic mustard provided more effective control than early weeding because late weeding allows second‐year plants to compete with garlic mustard seedlings for a longer period of time and (2) competition between first‐ and second‐year plants is responsible for alternating dominance of first‐year and second‐year garlic mustard plants.     

Escaping an evolutionary trap: preference and performance of a native insect on an exotic invasive host

Exotic plants may act as population sinks or evolutionary traps for native herbivores. The native butterfly Pieris oleracea lays eggs on garlic mustard, Alliaria petiolata, but larvae develop very poorly on this exotic invasive plant. We examined oviposition preference of individual females and larval performance of their offspring for individuals from one area where garlic mustard is well established and one where it is absent. These data were used to assess whether garlic mustard is being incorporated into or excluded from the diet. Females from the area without garlic mustard showed a wide range of preference, families had low larval survival on garlic mustard, and larval survivorship showed no correlation with mothers’ preferences. Females from the area with garlic mustard preferred it to the native host, and larval survivorship on garlic mustard was positively correlated with the mother’s preference. Individuals surviving on garlic mustard took longer to pupate and weighed >30% less compared to pupae reared on normal hosts. Our results suggest that where garlic mustard is well established P. oleracea may be adapting to this plant by both improved larval performance and increased adult female oviposition preference for it.     

Large plant size counteracts early seed predation during the extended flowering season of a Silene uniflora (Caryophyllaceae) population

A population of the alvar race of the perennial herb, Silene uniflora (Caryophyllaceae), which grows on thin soils in open alvar habitats on the Baltic island of Oland SE Sweden, was found to have an extended and more or less bimodal flowering phenology Large individuals produced flowers during both periods, while small, and presumably young, individuals only produced flowers in either of the two periods, or in part of both In the early flowering period plants were heavily infested by the seed-predatory larvae of a noctuid moth, Hadena confusa , but in the late flowering period only a small proportion of the fruits was attacked by the seed predators The proportion of flowers developing into fruits was consistently high throughout the season Both the number of seeds per capsule and the mass of seeds decreased over the flowering season However, the germination success of early and late seeds did not differ Thus, although differing in number of seeds, both early and late flowers contributed to individual reproductive success Large individuals started to flower early in the season and despite their high loss of seeds in the early part of the summer they contributed a larger number of seeds to the seed pool than smaller and later-flowering individuals Although selection was acting to favour later flowering during a year with high early seed predation, consistency of date of flower initiation and of relative predation impact on individuals across years was low suggesting that recurrent selection by seed predators is weak Seed predation, although heavy, is therefore judged to be unlikely to cause a significant evolutionary response on flowering phenology in this plant     

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.     

Growth, survival and generative reproduction in a population of a widespread annual hemiparasite Melampyrum pratense

Melampyrum pratense is one of the most successful root-hemiparasitic species of temperate Europe occurring in various habitats including both forest and open communities. The species shares its life history traits (large seeds, annuality, lack of clonality) with most of other hemiparasitic Orobanchaceae. Due to lack of vegetative propagation, the reproductive success of a M. pratense individual largely depends on the seed production. We used an individual-based observation of ontogenetic development of plants and analysed the development of spatial distribution of individuals to reveal links between fecundity of individuals and their vegetative growth in the context of population dynamics within one growing season. We demonstrated a tight dependency of seed production on vegetative growth and survival till the end of the growing season. Plants that were still alive and big (due to a high number of lateral branches) in the end of August produced the highest numbers of seeds. These were several times higher than the population average due to positively skewed distribution of seed production across individuals. The rate of premature mortality was rather low (below 50%) once seedlings successfully emerged and was most likely caused by intraspecific competition. By contrast, moderate level of herbivory (stem clipping by grazers) had a limited effect on the survival and the inflicted biomass losses seemed compensated. Therefore, despite being an annual, M. pratense appears well-adapted to its life in perennial-dominated forest communities, which is also underpinned by its hemiparasitic strategy providing essential resources and allowing to avoid below-ground competition.     

Impact of simulated herbivory on Alliaria petiolata survival, growth, and reproduction

In weed biological control, insect damage to target weeds can be simulated in invaded habitats to study potential responses of the plant to introduced natural enemies. In the present study, we investigated the impact of two levels of manual flower-shoot damage (shoots cut at tip or base) on Alliaria petiolata (garlic mustard) survival, size, and reproduction. Experiments were conducted in 2002 and 2003 using invasive field populations of A. petiolata under naturally varying plant densities. Plant survival was recorded, and size and reproduction parameters were measured. Manual flower-shoot damage had a significant effect on plant survival. In both years, fewer plants survived in the basal-cut treatment than in either the control (un-cut) or tip-cut treatment. Plant size and reproductive output were likewise reduced in the basal-cut treatment. In both years, total seed production was significantly lower in the basal-cut treatment than either the control or tip-cut treatment. When combined, increased mortality and reduced seed production of basal-cut plants greatly reduced the contribution these plants made to the seed bank. Plant density did not affect reproduction or plant size. The impacts of cutting were consistent across years and sites with distinct biotic and abiotic conditions, and A. petiolata densities. We anticipate that herbivore damage to A. petiolata populations by introduced biological control agents will likewise remain consistent under varying biotic and abiotic conditions if the agents are equally adapted to these.     

热带雨林林内小生境对木奶果和染木种子萌发与幼苗存活的影响

森林生态系统中小生境的差异可能影响种子萌发与幼苗的建立,并形成物种不同的分布格局。为探讨生境异质性对植物分布的影响,该研究在西双版纳热带季节性雨林中选取5个具有不同海拔和地形的小生境,人工散布木奶果与染木种子,定期监测种子萌发与幼苗存活,同步记录小生境的近地面（距地面3 cm）温度、近地面(3 cm)空气湿度和表层土壤(5 cm)含水量变化,并在实验室开展种子脱水和不同水势条件下的种子萌发实验。结果表明：（1）实验所选的5个小生境中,沟底地带的土壤水分含量在一年中始终都高于其他小生境;5个小生境的空气相对湿度在雨季与雾凉季都接近饱和,只有在干热季出现明显的差别,即随海拔的升高,空气湿度下降。同时,在旱季,近地面温度随海拔的升高而升高, 出现1~2个月的山地逆温现象。（2）木奶果与染木具有不同的种子传播与萌发规律,而且染木种子在不同生境下,其种子萌发率和幼苗存活率有显著的差异。其中：木奶果种子在雨季成熟散布并快速萌发,以幼苗的形式度过干热季,其种子可以在相对较低的水势下萌发;染木种子在雨季末期成熟并进入休眠期,以种子的形式度过干热季,其种子相对较抗脱水。研究认为,地形是决定西双版纳热带季节性雨林生境异质性的重要因子,小生境在温度和水分方面的差异在干热季时尤为明显,并对木奶果和染木种子的萌发或幼苗存活产生了重要影响, 而木奶果与染木种子自身的特性使其对于西双版纳季节性的干旱具有不同的适应对策。     

Impact of invasions by alien plants on soil seed bank communities: Emerging patterns

Much of our current understanding of the impact of invasive species on plant communities is based on patterns occurring in the above-ground vegetation, while only few studies have examined changes in soil seed banks associated with plant invasions, despite their important role as determinants of vegetation dynamics. Here, we reviewed the literature on the impact of plant invasions on the seed bank and we provide a quantitative synthesis using a meta-analysis approach. Specifically, (1) we quantified the impact of 18 invasive alien plants on (i) species richness and (ii) density of the seed banks of invaded communities, based on 58 pair-wise invaded-uninvaded comparisons (cases); we identified (2) the invasive taxa that are responsible for the largest changes in the seed bank; and (3) the habitats where substantial changes occur. Our study showed three major findings: (1) species richness (68% of cases) and density (58% of cases) were significantly lower in native seed banks invaded by alien plants; (2) species richness and density of native and alien species were remarkably lower in seed banks invaded by large, perennial herbs compared to uninvaded sites; and (3) invaded seed banks were often associated with a larger richness and/or abundance of alien species. This study indicates a need for additional seed bank data in invasion ecology to characterize species-specific and habitat-specific impacts of plant invasions, and to determine whether changes in the seed banks of native and alien species are a symptom of environmental degradation prior to a plant invasion or whether they are its direct result. The findings of this study help improve our capacity to predict the long-term implications of plant invasions, including limitations in the recruitment of native species from the seed bank and the potential for secondary invasions by seeds of other alien species.