Are Local Filters Blind to Provenance? Ant Seed Predation Suppresses Exotic Plants More than Natives

The question of whether species’ origins influence invasion outcomes has been a point of substantial debate in invasion ecology. Theoretically, colonization outcomes can be predicted based on how species’ traits interact with community filters, a process presumably blind to species’ origins. Yet, exotic plant introductions commonly result in monospecific plant densities not commonly seen in native assemblages, suggesting that exotic species may respond to community filters differently than natives. Here, we tested whether exotic and native species differed in their responses to a local community filter by examining how ant seed predation affected recruitment of eighteen native and exotic plant species in central Argentina. Ant seed predation proved to be an important local filter that strongly suppressed plant recruitment, but ants suppressed exotic recruitment far more than natives (89% of exotic species vs. 22% of natives). Seed size predicted ant impacts on recruitment independent of origins, with ant preference for smaller seeds resulting in smaller seeded plant species being heavily suppressed. The disproportionate effects of provenance arose because exotics had generally smaller seeds than natives. Exotics also exhibited greater emergence and earlier peak emergence than natives in the absence of ants. However, when ants had access to seeds, these potential advantages of exotics were negated due to the filtering bias against exotics. The differences in traits we observed between exotics and natives suggest that higher-order introduction filters or regional processes preselected for certain exotic traits that then interacted with the local seed predation filter. Our results suggest that the interactions between local filters and species traits can predict invasion outcomes, but understanding the role of provenance will require quantifying filtering processes at multiple hierarchical scales and evaluating interactions between filters.     

When do herbivores affect plant invasion? Evidence for the natural enemies and biotic resistance hypotheses

Two venerable hypotheses, widely cited as explanations for either the success or failure of introduced species in recipient communities, are the natural enemies hypothesis and the biotic resistance hypothesis. The natural enemies hypothesis posits that introduced organisms spread rapidly because they are liberated from their co‐evolved predators, pathogens and herbivores. The biotic resistance hypothesis asserts that introduced species often fail to invade communities because strong biotic interactions with native species hinder their establishment and spread. We reviewed the evidence for both of these hypotheses as they relate to the importance of non‐domesticated herbivores in affecting the success or failure of plant invasion.
To evaluate the natural enemies hypothesis, one must determine how commonly native herbivores have population‐level impacts on native plants. If native herbivores seldom limit native plant abundance, then there is little reason to think that introduced plants benefit from escape from these enemies. Studies of native herbivore‐native plant interactions reveal that plant life‐history greatly mediates the strength with which specialist herbivores suppress plant abundance. Relatively short‐lived plants that rely on current seed production for regeneration are most vulnerable to herbivory that reduces seed production. As such, these plants may gain the greatest advantage from escaping their specialist enemies in recipient communities. In contrast, native plants that are long lived or that possess long‐lived seedbanks may not be kept “in check” by native herbivores. For these species, escape from native enemies may have little to do with their success as exotics; they are abundant both where they are native and introduced.
Evidence for native herbivores providing biotic resistance to invasion by exotics is conflicting. Our review reveals that: 1) introduced plants can attract a diverse assemblage of native herbivores and that 2) native herbivores can reduce introduced plant growth, seed set and survival. However, the generality of these impacts is unclear, and evidence that herbivory actually limits or reduces introduced plant spread is scarce. The degree to which native herbivores provide biotic resistance to either exotic plant establishment or spread may be greatly determined by their functional and numerical responses to exotic plants, which we know little about. Generalist herbivores, through their direct effects on seed dispersal and their indirect effects in altering the outcome of native–non‐native plant competitive interactions, may have more of a facilitative than negative effect on exotic plant abundance.     

Ant interactions with native and exotic seeds in the Patagonian steppe: Influence of seed traits,disturbance levels and ant assemblage

Invasive plants may establish strong interactions with species in their new range which could limit or enhance their establishment and spread. These interactions depend upon traits of the invader and the recipient community, and may alter interactions among native species. In the Patagonian steppe we studied interactions of native ant assemblages with seeds of native and exotic plants, and asked whether ant–seed interactions differ with seed types and disturbance levels and whether the amount and type of ant–seed interactions can be predicted if both ant and seed traits are known. To characterize and quantify ant–seed interactions, we offered baits with large seeds of Pappostipa speciosa (native) and medium-sized elaiosome-bearing seeds of Carduus thoermeri (exotic), near and far from a road (high vs. low disturbed areas), and compared ant abundance and composition between areas. Interaction frequency was the highest for C. thoermeri seeds far from the road. Composition of ants interacting with C. thoermeri in these areas differed from that near the road and from that interacting with native seeds. Ant composition and abundance were similar between areas, but some species interacted more with exotic seeds in low disturbed areas. Ant foraging type predicted ant–seed interactions since the abundance of seed harvesters was positively correlated to interactions with P. speciosa, and that of generalists and predators, with interactions with C. thoermeri. The high interaction of ants with exotic seeds in low invaded areas suggests that ant activity could influence plant invasion, either by predating or dispersing seeds of invasive plants.     

Seed predation of two alien Opuntia species invading Mediterranean communities

Invasibility depends on the interaction of the introduced species with the abiotic and biotic factors of the recipient community. In particular, the biotic resistance posed by native herbivores has been claimed to be of great importance in controlling plant invasion. We investigated fruit and seed predation of two exotic Opuntia species within and between Mediterranean communities in order to determine how patterns of predation matched patterns of invasion. Predators were small mammals, presumably mice, which could consume more than 50% of the seeds produced. Predators could be equally effective in consuming fruit and single seeds. O. maxima fruits were slightly preferred to O. stricta fruits, but predators did not distinguish between seeds. Seed predation was more intense in invaded than in non-invaded communities. However, there was a high spatial variation in seed predation that did not always match patterns of invasion, suggesting that seed predation alone is not a good predictor of community invasibility to Opuntia. According to these results invasibility to Opuntia is limited in some (but not all) communities by native mice. Seed losses by predation were high for both species. However, we estimated that more than 75% of seeds dispersed by birds to non-invaded areas are not predated.     

Leaf-cutting ant nests near roads increase fitness of exotic plant species in natural protected areas

Understanding the mechanisms that promote the invasion of natural protected areas by exotic plants is a central concern for ecology. We demonstrated that nests of the leaf-cutting ant, Acromyrmex lobicornis, near roadsides promote the abundance, growth and reproduction of two exotic plant species, Carduus nutans and Onopordum acanthium, in a national park in northern Patagonia, Argentina and determine the mechanisms that produce these effects. Refuse dumps (RDs) from ant nests have a higher nutrient content than nearby non-nest soils (NNSs); foliar nutrient content and their 15N isotopic signature strongly suggest that plants reach and use these nutrients. Both species of exotic plants in RDs were 50-600% more abundant; seedlings had 100-1000% more foliar area and root and leaf biomass; and adult plants produced 100-300% more seeds than nearby NNS plants. Plants can thus gain access to and benefit from the nutrient content of ant RD, supporting the hypotheses that enhanced resource availability promotes exotic plant performance that could increase the likelihood of biological invasions. The two exotics produce an estimated of 8385000 more seeds ha(-1) in areas with ant nests compared with areas without; this exceptional increase in seed production represents a potential threat to nearby non-invaded communities. We propose several management strategies to mitigate this threat. Removal efforts of exotics should be focused on ant RDs, where plants are denser and represent a higher source of propagules.     

Seed predation of non-native species along a precipitation gradient

Generalist seed predators are an important factor shaping non-native species invasion. Their effect is highly influenced by abiotic conditions, yet how the importance of this biotic filter changes in a gradient of abiotic conditions is still poorly understood. In this study, we assessed seed predation of non-native conifer species along a precipitation gradient in north-western Argentinean Patagonia. We performed a seed removal experiment over a 15-day period during the fall in sites where annual precipitation ranges from 600 to 1600 mm and vegetation ranges from steppe to forest. We used the three most common forestry species in the study area, including two species known to invade areas adjacent to plantations and one native species. Seed predation was higher in sites with higher precipitation, but no significant differences in seed predation were found among species. Small mammals were the dominant group predating seeds on this gradient. These findings highlight the importance of abiotic conditions mediating plant-granivore interactions, and the influence that abiotic conditions may have on biotic resistance to invasion.     

Integration of exotic seeds into an Azorean seed dispersal network

Seed dispersal plays a central role in plant ecology. Among animals, birds are particularly important seed dispersers, often incorporating exotic plants into their diets and facilitating their integration in the communities. Network theory offers a highly informative framework to study the structural and functional attributes of complex interactions networks. We used information from bird fecal samples to build a quantitative seed dispersal network for the last fragment of native laurel forest in the island of São Miguel—Azores with three specific objectives: (1) to assess the integration of exotic seeds into seed dispersal; (2) to evaluate the impact of exotic plants in network structure; (3) to test the potential of an exotic species to reduce the seed dispersal of a co-occurring native, via competition for seed dispersers. The seed dispersal network was based on the analysis of 1,121 droppings and described 74 unique interactions between 41 plant species and 7 bird species. Exotic seeds deeply infiltrated into the seed dispersal network forming the majority (59 %) of seeds in the droppings and including those of three globally invasive plants. Overall, birds depended equally on native and exotic fruits despite the lower abundance of the latter in the study area. In an experiment, birds did not show a preference for fruits of the exotic Leycesteria formosa over the native Vaccinium cylindraceum consuming them equally. However, the presence of the exotic plant negatively affected the number of native seeds dispersed, by diverting some of the consumers of the native fruits. Taken altogether the results reveal an alarming invasion level of seed dispersal systems in one of the last remnant native forests of the Azores.     

Can a breakdown in competition–colonization tradeoffs help explain the success of exotic species in the California flora?

Determining combinations of functional traits that allow a species to colonize new habitats has been central in the development of invasion ecology. Species able to establish in new communities harbor abilities or traits that allow them to use resources or tolerate stress in ways that native species cannot. Tradeoffs among species functional traits along the competition–colonization (CC) continuum, where competitive ability is a decreasing function of dispersal capacity, may allow invasive species to establish themselves in new habitats. The California flora offers a well‐characterized model system to examine whether native and exotic species differ in the distribution of functional traits and to examine whether a breakdown of the CC tradeoff is present. We used a random subset of 1000 plants and examined seed traits and life form characteristics along with their seed size and adult height using the Jepson Manual of the plants of California. To test the hypothesis that active dispersal strategies aid in the success of exotic species, we classified species into four seed types according to the presence/absence of mechanisms associated with efficient dispersal. In addition, for each species we compiled data on seed size and adult plant height. We conducted all comparisons between native and exotic species within the four most speciose families to control for potential taxonomic non‐independence. Exotic species had smaller seed size but greater plant height than natives of the same families. On the other hand, exotic species also displayed significantly greater proportions of functional traits that enhanced dispersal ability. Additionally, certain sets of functional traits were significantly associated with exotic species, such as annual life histories with small seeds and high dispersal capacity. In the random subset of the California flora examined, exotics of the most speciose plant families show functional trait combinations that appear to violate the tradeoff structures observed in their California counterparts. Our results suggest that taxonomically controlled comparisons of the CC tradeoff structure between natives and exotic species may shed light of the capacity of those exotic species invasive ability to colonize new habitats.     

Effects of an exotic plant and habitat disturbance on pollinator visitation and reproduction in a boreal forest herb

The invasion of exotic species into natural habitats is considered to be a major threat to biodiversity, and many studies have examined how exotic plants directly affect native plant species through competitive interactions for abiotic resources. However, although exotics can have potentially great ecological and evolutionary consequences, very few researchers have studied the effect of exotics on the interactions between plants and their mutualistic partners, such as pollinators, and none have reported on such impacts in logged and undisturbed boreal forest ecosystems. Here we show how experimental introductions of an exotic plant species (Phacelia tanacetifolia Bentham) affect pollinator visitation and female reproductive success of a native plant (Melampyrum pratense L.) in recently disturbed (i.e., logged) and in undisturbed boreal forest habitats. The presence of Phacelia significantly increased the number of bumble bees entering plots in both habitat types. However, the exotic species had a strong negative impact on the visitation rate to the native species in both habitat types. Despite this negative impact on pollinator visitation, the exotic had no effect on female reproductive success of the native species in any habitat. Our results show that seed production may be more robust than pollinator visitation to exotic invasion, irrespective of habitat disturbance history.     

Is <Emphasis Type="Italic">Pinus radiata</Emphasis> invading the native vegetation in central Chile? Demographic responses in a fragmented forest

Forest fragmentation facilitates the invasion of exotic species. This threat may be especially severe if forest fragments are surrounded by plantations of exotic species like Pinus radiata, an aggressive colonizer and shade-intolerant tree that has invaded successfully several native ecosystems of the southern hemisphere. In this study, we experimentally tested if the conditions of a successful seedling establishment P. radiata are fulfilled at the Coastal Maulino forest, an endemic fragmented forest of central Chile. Results demonstrated that seeds are dispersed into the native forests, however seedling establishment occurs only at the edges. We conclude that this exotic species is not invading native forests up to date. However, we suggest to conduct evaluations of seed rain and seedling establishment in the long term, in order to monitor the fate of this exotic species in fragmented native forest of Central Chile.     

Enemy release or invasional meltdown? Deer preference for exotic and native trees on Isla Victoria,Argentina

Abstract How interactions between exotic species affect invasion impact is a fundamental issue on both theoretical and applied grounds. Exotics can facilitate establishment and invasion of other exotics (invasional meltdown) or they can restrict them by re‐establishing natural population control (as predicted by the enemy‐release hypothesis). We studied forest invasion on an Argentinean island where 43 species of Pinaceae, including 60% of the world's recorded invasive Pinaceae, were introduced c. 1920 but where few species are colonizing pristine areas. In this area two species of Palearctic deer, natural enemies of most Pinaceae, were introduced 80 years ago. Expecting deer to help to control the exotics, we conducted a cafeteria experiment to assess deer preferences among the two dominant native species (a conifer, Austrocedrus chilensis, and a broadleaf, Nothofagus dombeyi) and two widely introduced exotic tree species (Pseudotsuga menziesii and Pinus ponderosa). Deer browsed much more intensively on native species than on exotic conifers, in terms of number of individuals attacked and degree of browsing. Deer preference for natives could potentially facilitate invasion by exotic pines. However, we hypothesize that the low rates of invasion currently observed can result at least partly from high densities of exotic deer, which, despite their preference for natives, can prevent establishment of both native and exotic trees. Other factors, not mutually exclusive, could produce the observed pattern. Our results underscore the difficulty of predicting how one introduced species will effect impact of another one.     

Native granivores reduce the establishment of native grasses but not invasive <Emphasis Type="Italic">Bromus tectorum</Emphasis>

Seed predation can structure plant communities by imposing strong population controls on some species but not others. In this context, studies from various ecosystems report that native granivores selectively forage for seeds from native species over seeds from exotic invaders, which could disproportionately favor the establishment of invaders and facilitate their dominance in communities. However, few studies have connected selective foraging for native seeds to differential patterns of establishment among native and invasive species. Thus, the extent to which preferential foraging for native seeds favors the establishment of invasive plants is unclear. Here, we used experimental seed additions and exclosure treatments at five field sites distributed across?≈?80,000 km² of the Great Basin Desert, USA to compare the effects of rodent foraging on the establishment of less-preferred cheatgrass (Bromus tectorum—an annual species native to Eurasia that is exotic and highly invasive across the Great Basin) and four species of more-preferred native grasses that commonly co-occur with cheatgrass. Rodent foraging reduced the establishment of each native species by at least 80% but had no effect on the establishment of cheatgrass, and this finding was consistent across study sites. Our results suggest that selective foraging for native species may favor the establishment of cheatgrass over native grasses, potentially exacerbating one of the most extensive plant invasions in North America.     

Deposition of exotic bird-dispersed seeds into three habitats of a fragmented landscape in the northeastern United States

Dispersal by frugivorous birds facilitates invasion by many exotic plants. We measured the seed rain of ornithochorous native and exotic plants at three habitats of a fragmented landscape of the northeastern United States for 1 year. We studied maple-beech forests, old fields, and abandoned conifer plantations. Across all sites we collected 2,196 ornithochorous seeds, including seeds from six exotic species and 10 native species. The majority (90%) of collected seeds were from exotic species. Seed dispersal was broadly similar among habitats, though seed rain of exotic species was higher in old fields than forested habitats. Seed rain was not strongly influenced by artificial perches for most species. However, seeds of exotic species were more commonly found in traps under an artificial perch in old fields. Seed rains for the exotic Elaeagnus umbellata, Rhamnus cathartica, and Rosa multiflora were positively associated with local density of mature plants. Seed rain of R. cathartica was positively associated with abundance of seedlings but not saplings, suggesting that post-dispersal mortality was important. Seed dispersal of the exotic Lonicera spp. was high in all habitats, accounting for 66% of all seeds collected. With the exception of Lonicera spp., seed rain of common exotic invaders was affected by the abundance of seed sources, and these species might be effectively controlled by elimination of local fruiting plants. Fruits of Lonicera morrowii, which has extensively invaded our area, are apparently a common component in the diet of frugivores.     

Native Consumers and Seed Limitation Constrain the Restoration of a Native Perennial Grass in Exotic Habitats

Native consumers and seed limitation may be particularly important in the restoration of native plants where they have been displaced by exotic plants. We used experimental exclosures and seed additions to examine the role of native mammalian consumers and seeding density (500 or 1,000 seeds/m²) in affecting the establishment of a native perennial grass, Purple needlegrass ( Nassella pulchra ), in the grasslands of California. To focus solely on consumers and propagule density, experimental areas were tilled and weeded. Consumers were important determinants of restoration success: averaged across propagule density, consumers reduced N. pulchra seedling recruitment by nearly 30%, reduced seedling height by 44%, reduced plant establishment by 52%, and reduced reproductive tiller length by 43%. Small rodents affected seedling establishment, especially where seeding density was high but did not affect seedling height. Plots accessible by squirrels and rabbits exhibited significantly decreased seedling height and plant establishment, whereas there was no additional impact of allowing large consumers (i.e., deer) access. Despite strong, spatially variable effects of consumers, doubling seed density led to nearly doubled N. pulchra establishment on average. Consumer effects were persistent, shaping N. pulchra abundance in the subsequent growing season and remaining evident over 18 months after the experiment was initiated. Our work suggests that, despite strong consumer effects, seed addition may be a viable strategy for restoration of N. pulchra in invaded areas where it has been displaced by exotic plants, especially when combined with restoration strategies that reduce competition with exotic plants.     

Responses of exotic plant species to fires in Pinus ponderosa forests in northern Arizona

Abstract. Changes in disturbance due to fire regime in southwestern Pinus ponderosa forests over the last century have led to dense forests that are threatened by widespread fire. It has been shown in other studies that a pulse of native, early‐seral opportunistic species typically follow such disturbance events. With the growing importance of exotic plants in local flora, however, these exotics often fill this opportunistic role in recovery. We report the effects of fire severity on exotic plant species following three widespread fires of 1996 in northern Arizona P. ponderosa forests. Species richness and abundance of all vascular plant species, including exotics, were higher in burned than nearby unburned areas. Exotic species were far more important, in terms of cover, where fire severity was highest. Species present after wildfires include those of the pre‐disturbed forest and new species that could not be predicted from above‐ground flora of nearby unburned forests.     

A null model of exotic plant diversity tested with exotic and native species-area relationships

At large spatial scales, exotic and native plant diversity exhibit a strong positive relationship. This may occur because exotic and native species respond similarly to processes that influence diversity over large geographical areas. To test this hypothesis, we compared exotic and native species–area relationships within six North American ecoregions. We predicted and found that within ecoregions the ratio of exotic to native species richness remains constant with increasing area. Furthermore, we predicted that areas with more native species than predicted by the species–area relationship would have proportionally more exotics as well. We did find that these exotic and native deviations were highly correlated, but areas that were good (or bad) for native plants were even better (or worse) for exotics. Similar processes appear to influence exotic and native plant diversity but the degree of this influence may differ with site quality.     

外来植物入侵的化感作用机制探讨

在最近的几十年,在全球范围内的外来植物入侵的频率达到了前所未有的水平,所以对其入侵机制的研究显得尤为重要。目前,已有几种假说从不同的侧面解释外来植物的成功入侵。本文在综述前人工作基础上,从化感作用的角度解释外来植物入侵,并给出解释模式。由于本土动植物对某些外来植物释放的化感物质比较敏感,外来植物可以利用它的“化学武器”与本土植物竞争,抵御植食性动物的取食和病源菌的感染,从而在与本土物种的相互干扰中占据优势,实现成功入侵。进一步从理论和应用的角度探讨了化感作用作为外来植物入侵的机制给我们的启示,并对今后这方面研究提出几点建议和看法。     

Characterizing the microhabitats of exotic species in Illinois shale barrens

Like native species, exotics form part of the mosaic structure of plant communities. However, their role in these communities is unclear. Thus, we ask, are invading exotic species present in recognizable microhabitats within plant communities? We compared the microhabitat of exotic species and random locations to determine if exotic plants occupied a recognizable subset of habitats within three barren communities. Fourteen exotic species were identified. The factors that characterized their habitats varied between barrens and among species. Considered as a group, exotic microhabitats comprised a subset of available habitats within the barrens, specifically, edge habitats rather than the open conditions of the barrens interior. Soil properties such as low temperature or neutral pH, and high litter or woody species cover, were common factors associated with the occurrence of exotics. Individual exotic species occupied different habitats within those identified for exotics as a group, but the range of variation between the different species was not generally significant. Within each barren, exotics occupied a distinct subset of microhabitats. This distribution suggests that exotics are not widely dispersed within these areas. Site conditions may be restricting the establishment of exotics to specific microhabitats, and in addition, some exotics may be altering the microhabitats that they colonize. Therefore, management efforts toward maintaining open conditions in the barrens may also discourage further encroachment by exotics into these areas.     

Diminishing importance of elaiosomes for acacia seed removal in non-native ranges

Myrmecochorous plants produce seeds with lipid-rich appendages (elaiosomes) which act as a reward for seed-dispersing ants. Seed dispersal is important for exotic species, which often need to establish new mutualistic interactions in order to colonize new non-native habitats. However, little is known about the importance of elaiosomes for seed removal in many of their non-native ranges. We studied ant–seed interactions of elaiosome-bearing and elaiosome-removed seeds of the Australian trees Acacia dealbata and Acacia longifolia in order to assess the relative importance of elaiosomes for seed removal between their native (Australia) and non-native (Portugal) ranges. In Portugal, we also studied the co-occurring native plant species with myrmecochorous seeds, Pterospartum tridentatum and Ulex europaeus, across three contiguous levels of acacia invasion: control (i.e. no acacia), low, and high acacia tree density. Acacia seeds were successfully removed by ants in their non-native region by a diversified assemblage of ant species, even in sites where native plants interacted with only one specialized ant species. In the invaded range, diminishing relative importance of elaiosomes was associated with changes in the ant community due to acacia invasion, and for A. dealbata, ant species richness decreased with increasing acacia tree density. Although seed removal was high for both acacia species, the importance of elaiosomes was proportionally lower for A. dealbata in the non-native region. Native plant species experienced significant reductions in seed removal in areas highly invaded by acacia, identifying another mechanism of displacement of native plants by acacias.     

Processes regulating the invasion of European buckthorn (Rhamnus cathartica) in three habitats of the northeastern United States

Invasion by exotic plants often is restricted by processes, such as seed predation, acting on early life-history stages; however, the relative importance of these processes might vary among habitats. Modern human land use has created a mosaic of habitats in many landscapes, including the landscape of the northeastern United States. European buckthorn (Rhamnus cathartica) is an exotic plant that has achieved varying success in North American habitats. We studied dispersal, seed survival, germination, and seedling survival of buckthorn populations at four plots in each of sugar maple (Acer saccharum) forests, old fields, and abandoned conifer plantations in central New York State. Dispersal was low in maple forests, as evidenced by low collection rates of R. cathartica seeds in seed traps. Rates of post-dispersal seed predation were highest in maple forests and lowest in old fields, suggesting greater use of maple forests by granivorous rodents. Germination rates did not vary among seeds planted in soils of these habitats when studied in the laboratory despite differences in soil pH. Survival of transplanted seedlings was low in maple forests relative to old fields and plantations. Buckthorn invasion of old fields and abandoned plantations was not strongly constrained by factors we considered, and the buckthorn populations in these habitats were large. A combination of low dispersal by frugivores, low seed survival due to predation, and low seedling survival due to dim light conditions apparently prevents R. cathartica from invasion of intact maple forests of our area. Native fauna and canopy closure may act synergistically to reduce success of invasive plants in natural habitats.