Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests

Species interactions and their indirect effects on the availability and distribution of resources have been considered strong determinants of community structure in many different ecological systems. In deciduous forests, the presence of overstory trees and shrubs creates a shifting mosaic of resources for understory plants, with implications for their distribution and abundance. Determination of the ultimate resource constraints on understory vegetation may aid management of these systems that have become increasingly susceptible to invasions by non-native plants. Microstegium vimineum (Japanese grass) is an invasive annual grass that has spread rapidly throughout the understory of forests across the eastern United States since it was first observed in Tennessee in 1919. M. vimineum occurs as extensive, dense patches in the understory of eastern deciduous forests, yet these patches often exhibit sharp boundaries and distinct gaps in cover. One example of this distributional pattern was observed relative to the native midstory tree Asimina triloba (pawpaw), whereby dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy. We conducted field and greenhouse experiments to test several hypotheses regarding the causes of this observed pattern of M. vimineum distribution, including allelopathy, seed dispersal, light limitations, and soil moisture, texture, and nutrient content. We concluded that light reduction by the A. triloba canopy was the environmental constraint that prevented establishment of M. vimineum beneath this tree. Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer. It is likely that other midstory species influence the distribution and abundance of other herb-layer species, with implications for management of understory invasive plant species.     

Competitive context alters plant-soil feedback in an experimental woodland community

Recent findings on feedback between plants and soil microbial communities have improved our understanding of mechanisms underlying the success and consequences of invasions. However, additional studies to test for feedback in the presence and absence of interspecific competition, which may alter the strength or direction of feedbacks, are needed. We tested for soil microbial feedback in communities of the invasive grass Microstegium vimineum and commonly co-occurring native plant species. To incorporate competitive context, we used a factorial design with three plant treatments (M. vimineum alone, M. vimineum with the native plant community, and the native community without M. vimineum) and two soil inoculum treatments (experimentally invaded and uninvaded soil). When competing with M. vimineum, native communities were 27% more productive in invaded than uninvaded soil. In contrast, soil type did not significantly affect M. vimineum biomass or fecundity. At the community level, these results indicate a net negative soil microbial feedback when native plants and M. vimineum are grown in competitive mixture, but not when they are grown separately. Since positive, not negative, feedback is associated with dominance and invasion, our findings do not support plant–soil feedback as a driver of invasion in this species. Our results do show that the importance of soil feedback can change with competitive context. Such context-dependency implies that soil feedback may change when competitive interactions between natives and invading species shift as invasions progress.     

The Roles of Competition and Disturbance in a Marine Invasion

Two hypotheses for the decline of native species are the superior exploitation of disturbance by exotic species and the competitive displacement of native species by their exotic counterparts. Theory predicts that functional similarity will increase the intensity of competition between native and invasive species. Ecologically important “foundation” species, Zostera marina and other seagrasses have globally declined during the past century. This study used transplant and vegetation removal experiments to test the hypotheses that disturbance and competitive interactions with an invasive congener (Z. japonica) are contributing to the decline of native Z. marina in the northeastern Pacific. Interspecific competition reduced Z. marina and Z. japonica above-ground biomass by 44 and 96%, respectively, relative to intraspecific competition. Disturbance substantially enhanced Z. japonica productivity and fitness, and concomitantly decreased Z. marina performance, effects that persisted two years following substratum disturbance. These results demonstrate that disturbance and competitive interactions with Z. japonica reduce Z. marina performance, and suggest that Z. japonica’s success as an invasive species stems dually from its ability to persist in competition with Z. marina and its positive response to disturbance. These results highlight the importance of understanding the interconnected roles of species interactions and disturbance in the decline of seagrass habitats, and provide a rationale for amending conservation policy in Washington State. In the interest of conserving native eelgrass populations, the current policy of protecting both native and invasive Zostera spp. should be refined to differentiate between native and invader, and to rescind the protection of invasive eelgrass.     

Competitive relationships between two contrasting but coexisting grasses

Leymus chinensis (Trin.) Tzvelev and Phragmites communis Trin. are co-dominant grasses in both nutrient-poor and rich steppes in the Songnen Plains of northeastern China. A replacement series experiment was conducted along a five-level nutrient gradient to test for changes in the competitive relationships of the species with nutrient availability. There were significant effects of nutrient level and species proportion on mean plant height, biomass and rhizome length. Facilitation was found in these plant attributes for both species when grown in mixture compared with monoculture, especially at high nutrient levels. For example, L. chinensis grew taller and intercepted more light when in mixture. The aboveground dominance of P. communis did not decrease in mixture, possibly because it benefited from belowground interactions with L. chinensis. Biomass allocation was apparently influenced by both above and belowground competition. For example, L. chinensis allocated much more biomass to shoots and less to roots when neighbor number was increased, suggesting that aboveground competition occurred. Root: shoot ratios of L. chinensis in monoculture decreased gradually with increasing nutrient availability, whereas the ratio in mixture declined rapidly, implying intense competition for light at high levels. Our results support the hypothesis that nutrient competition is more important than light competition when soil nutrients are scarce. Although the interactions between these species benefit both, P. communis is more likely to displace L. chinensis at high nutrient availability.     

Influence of litter removal and mineral soil disturbance on the spread of an invasive grass in a Central Hardwood forest

Soil and litter disturbances within Central Hardwood forests may facilitate exotic plant species invasion of interior forest areas. Microstegium vimineum is an annual exotic grass that has become common throughout the Southeastern United States. Three replicates of three different mineral soil and litter disturbance treatments, plus a control with no disturbance, were established on the leading edge of M. vimineum patches prior to seed fall. All patches were located in areas with similar forest canopy structure and slope in three Central Hardwood forest stands prior to seed fall. At the beginning of the following growing season, each individual M. vimineum seedling was mapped within the treatment plots. The mean number of M. vimineum individuals that established within each treatment did not differ significantly from the control. The distance at which 90% of the individuals had spread, and the overall mean distance spread were significantly farther for the litter removal treatment than the control. The farthest individual seedling from the boundary of existing patches in both the litter removal and the mineral soil disturbance and litter removal treatments were significantly farther than the control. The individuals that spread the farthest are of most concern due to the large number of viable seed that a single M. vimineum plant can produce. These results suggest that disturbance of the litter layer may increase the spread rate of M. vimineum and facilitate its invasion of new habitats, and that leaving litter layers intact may slow the spread of M. vimineum.     

Local‐scale biotic interactions embedded in macroscale climate drivers suggest Eltonian noise hypothesis distribution patterns for an invasive grass

A hierarchical view of niche relations reconciles the scale‐dependent effects of abiotic and biotic processes on species distribution patterns and underlies most current approaches to distribution modeling. A key prediction of this framework is that the effects of biotic interactions will be averaged out at macroscales – an idea termed the Eltonian noise hypothesis (ENH). We test this prediction by quantifying regional variation in local abiotic and biotic niche relations and assess the role of macroclimate in structuring biotic interactions, using a non‐native invasive grass, Microstegium vimineum, in its introduced range. Consistent with hierarchical niche relations and the ENH, macroclimate structures local biotic interactions, while local abiotic relations are regionally conserved. Biotic interactions suppress M. vimineum in drier climates but have little effect in wetter climates. A similar approach could be used to identify the macroclimatic conditions under which biotic interactions affect the accuracy of local predictions of species distributions.     

The putative niche requirements and landscape dynamics of Microstegium vimineum: an invasive Asian grass

The theoretical foundations of population and community ecology stress the importance of identifying crucial niche requirements and life history stages of invasive species and, in doing so, give insight into research and management. We focus on Microstegium vimineum, an invasive grass which is causing marked changes in the structure and function of US forests. We describe M. vimineum’s life history and habitat characteristics, infer its niche requirements and synthesize this information in the context of population dynamics and management. Based on the results synthesized here, M. vimineum’s crucial niche requirements appear to be light (reproductive output), soil moisture (reproductive output, seedling recruitment) and aboveground coverage by leaf-litter and competing species (seedling recruitment and survival). These data suggest a source-sink dynamic might allow M. vimineum to disperse and thrive along sunny, and sometimes wet, edge habitats and, in turn, these populations might act as source populations for adjacent shady forest habitats. By evaluating M. vimineum in the context of its stage-specific requirements, we highlight potential weaknesses in its life history that provide strategies for effective management.     

Restoring restoration: removal of the invasive plant <Emphasis Type="Italic">Microstegium vimineum</Emphasis> from a North Carolina wetland

Restoration sites are vulnerable to plant invasions due to habitat and resource alteration. We conducted an invasive plant-removal study at a wetland restoration in the North Carolina Piedmont, a site dominated by the non-native invasive, Microstegium vimineum. Paired plots (M. vimineum hand-weeded and unweeded) were established and maintained to monitor response of plant species richness and diversity. Plots increased from 4 to 15 species m⁻² after three growing seasons of M. vimineum removal and 90% of the newly establishing species were native. Weeding ceased in the fourth growing season and M. vimineum rapidly re-invaded. Formerly weeded plots increased to 59% (±11% SE) M. vimineum cover, 25 of 51 plant species disappeared from the plots, and species richness decreased to an average of <8 species m⁻². Our results show that we can quickly establish an abundant, diverse community with invasive removal, but that persistent effort is required to monitor and maintain the long-term viability of this community.     

Unadapted behaviour of native,dominant ant species during the colonization of an aggressive,invasive ant

Among the factors driving the invasive success of non-indigenous species, the “escape opportunity” or “enemy release” hypothesis argues that an invader’s success may result partly from less resistance from the new competitors found in its introduced range. In this study, we examined competitive interactions between the little fire ant Wasmannia auropunctata (Roger) and ant species of the genus Pheidole in places where both are native (French Guiana) and in places where only species of Pheidole are native (New Caledonia). The experimental introduction of W. auropunctata at food resources monopolized by the Pheidole species induced the recruitment of major workers only for the Guianian Pheidole species, which were very effective at killing Wasmannia competitors. In contrast, an overall decrease in the number of Pheidole workers and no recruitment of major workers were observed for the New Caledonian species, although the latter were the only ones able to kill the Wasmannia workers. These results emphasize the inappropriate response of native dominant New Caledonian species to W. auropunctata and, thus, the importance of enemy recognition and specification in the organization of ant communities. This factor could explain how invasive animal species, particularly ants, may be able to successfully invade species-rich communities.     

Compensation and Plasticity in an Invasive Plant Species

Plant plasticity under varying resources is one character that contributes to competitiveness and invasibility. The plasticity of plant modules of the invasive Japanese stilt grass (Microstegium vimineum) was examined under different light and nutrient levels using a balanced field and greenhouse study. Ample light and nutrients resulted in the greatest biomass accumulation in all plant modules. However, M. vimineum showed extreme plasticity, producing stolons and flowers (reproductive structures) under a wide range of environmental conditions except infertile, low light. The addition of nutrients under various shading regimes compensated for lower light levels, rendering little change in the allocation of carbon to reproductive structures. Data from the field study corroborate the greenhouse results, but also suggest plasticity in response to competition.     

Differences in life history strategies between an invasive and a competing resident seed predator

Competitive interactions may arise from biological invasion if a successful invasive species requires the ecological niche of a resident one. Life-history traits that make a species a successful invader are of particular interest in elucidating both invasion success and how interspecific competition may emerge. In southeastern France, the invasion of cedar forests by the seed chalcid Megastigmus schimitscheki generated competitive relationships with the resident M. pinsapinis for the exploitation of the seed resource. Among the numerous ecological traits allowing these seed predators to exploit their niche successfully, the timing of adult emergence, initial egg load and age-specific realized fecundities of females were investigated to help understanding the issue of such interspecific relationships. Spring adult emergence of M. schimitscheki under natural conditions was significantly earlier than that of M. pinsapinis, suggesting that an advantage for the access to the seed resource for the invasive species may be associated with this trait. Initial egg load was significantly higher in M. schimitscheki than in M. pinsapinis and the analysis of age-specific realized fecundity in semi-natural conditions indicated that both M. schimitscheki and M. pinsapinis females lay a large proportion of their eggs during the early days of their lives. In the light of these findings, both earlier phenology and higher reproduction abilities of M. schimitscheki may have the potential to confer an advantage within a competitive context with M. pinsapinis through enhanced seed resource preemption. This may potentially explain the invasiveness of M. schimitscheki in southern France despite the presence of its closely related competitor M. pinsapinis.     

Invasion impacts local species turnover in a successional system

Exotic plant invasions are often associated with declines in diversity within invaded communities. However, few studies have examined the local community dynamics underlying these impacts. Changes in species richness associated with plant invasions must occur through local changes in extinction and/or colonization rates within the community. We used long‐term, permanent plot data to evaluate the impacts of the exotic vine Lonicera japonica. Over time, species richness declined with increasing L. japonica cover. L. japonica reduced local colonization rates but had no effect on extinction rates. Furthermore, we detected significant reductions in the immigration of individual species as invasion severity increased, showing that some species are more susceptible to invasion than others. These findings suggest that declines in species richness associated with L. japonica invasion resulted from effects on local colonization rates only and not through the competitive displacement of established species.     

Is the increased vigour of invasive weeds explained by a trade-off between growth and herbivore resistance?

Blossey and Nötzold (1995) recently hypothesised that the increased vigour of certain invasive plant species has been at the expense of defences against natural enemies. A prediction of their evolution of increased competitive ability (EICA) hypothesis is that invasive genotypes are relatively poorly defended. We tested this prediction with herbivore bioassays and with direct quantification of plant secondary metabolites comparing non-indigenous genotypes of Lythrum salicaria L. (purple loosestrife) with indigenous forms. The herbivore bioassays revealed no significant intra-specific variation in herbivore resistance between indigenous and non-indigenous hosts. The phenolic content of L. salicaria leaves was significantly higher in indigenous genotypes, as predicted by the EICA hypothesis. The average phenolic content of leaves (regardless of their origin) was, however, low, implying that the role of plant phenolics in purple loosestrife anti-herbivore defence is probably limited. It is suggested that the EICA hypothesis, as tested in the current study, does not explain the increased vigour of L. salicaria in non-indigenous habitats.     

Competition and coexistence of rhizomatous perennial plants along a nutrient gradient

I studied competition and coexistence of three tall clonal perennial plant species, Calamagrostis epigejos (L.) Roth, Solidago canadensis L., and Tanacetum vulgare L. along a gradient of soil productivity over five years. A replacement series field experiment was conducted with high, moderate and low fertility levels in 1m×1m plots. There were significant effects of soil type on ramet density (P<0.001), mean height (P<0.01), and total biomass (P<0.01). Ramet density, mean height, and total biomass increased with increasing soil fertility. There were also significant effects of mixture on ramet density (P<0.01), but not on mean height and total biomass for all species. Significant neighbor effects on ramet density and total biomass (P<0.01) were found for Solidago, showing that it is important whether Tanacetum or Calamagrostis is its neighbor within mixtures. During the five years there was only one case of competitive exclusion: Calamagrostis excluded Solidago on the most fertile substrate in the fifth growing season. In most cases species coexisted over the five years. Each of the three species was able to dominate in at least one combination of substrate type and mixture. The experiment showed that asymmetric competition for light on substrates of high fertility, symmetric competition for nutrients on nutrient-poor soil and positive interactions especially on substrates of intermediate fertility played a role. A founder effect was evident in aggregated mixtures of Calamagrostis and Solidago on the nutrient-rich substrate. A conceptual model of the relative importance of root competition for soil nutrients, shoot competition for light, and positive interactions along the fertility gradient is presented. The model emphasizes that positive interactions play an important role over a broad range of the productivity scale with a peak at intermediate levels of fertility. On the substrate of high productivity shoot competition for light is more important than positive interactions and root competition for soil nutrients as well. The competitive superiority of Calamagrostis on the most productive substrate was evident only in the long run. Rare events like extreme summer drought or selective herbivore pressure caused a switch in dominance in mixtures with Solidago, respectively Tanacetum. The guerrilla growth strategy of Calamagrostis and interference competition through a dense cover of aboveground biomass and litter could further cause competitive exclusion.     

Zooplankton seasonal dynamics in a recently filled mine pit lake: the effect of non-indigenous <Emphasis Type="Italic">Daphnia</Emphasis> establishment

We examined the temporal and vertical dynamics of zooplankton in Weavers Lake, New Zealand, between October 2004 and October 2005, at a time when it was colonised by a non-indigenous Daphnia species. Zooplankton community composition changed during the study from one of rotifer dominance (e.g. Asplanchna, Polyarthra, Brachionus and Keratella species) to cladoceran (Daphnia dentifera) dominance. Temporal changes in zooplankton community composition were strongly associated with a gradual increase in lake water clarity, and were attributable to the highly efficient filter feeding of D. dentifera. The corresponding reduction in rotifer densities may have resulted from the superior competitive abilities of the newly established Daphnia. As Daphnia were rare inhabitants of New Zealand lakes before 1990, the arrival and rapid spread of the non-indigenous D. dentifera has lead to widespread changes in both water clarity and zooplankton community composition. An apparent lack of mixing in the lake was facilitated by the lake’s extremely small surface area:depth ratio. However, we conclude that physical features of the lake had minimal influence on water clarity relative to the invasion of D. dentifera.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Two new species of <Emphasis Type="Italic">Lirula</Emphasis> on <Emphasis Type="Italic">Abies</Emphasis> from Japan

Two new species of Lirula (L. japonica and L. exigua) on Abies mariesii collected in subalpine areas of northern Japan are described as members of Rhytismatales, Discomycetes. Lirula japonica causes needle cast in fir, but L. exigua seems to occur on the needles of physically damaged twigs. Morphological characteristics of both species are discussed.     

Nitrogen uptake and preference in a forest understory following invasion by an exotic grass

Plant–soil interactions have been proposed as a causative mechanism explaining how invasive plant species impact ecosystem processes. We evaluate whether an invasive plant influences plant and soil-microbe acquisition of nitrogen to elucidate the mechanistic pathways by which invaders might alter N availability. Using a ¹⁵N tracer, we quantify differences in nitrogen uptake and allocation in communities with and without Microstegium vimineum, a shade-tolerant, C₄ grass that is rapidly invading the understories of eastern US deciduous forests. We further investigate if plants or the microbial biomass exhibit preferences for certain nitrogen forms (glycine, nitrate, and ammonium) to gain insight into nitrogen partitioning in invaded communities. Understory native plants and M. vimineum took up similar amounts of added nitrogen but allocated it differently, with native plants allocating primarily to roots and M. vimineum allocating most nitrogen to shoots. Plant nitrogen uptake was higher in invaded communities due primarily to the increase in understory biomass when M. vimineum was present, but for the microbial biomass, nitrogen uptake did not vary with invasion status. This translated to a significant reduction (P < 0.001) in the ratio of microbial biomass to plant biomass nitrogen uptake, which suggests that, although the demand for nitrogen has intensified, microbes continue to be effective nitrogen competitors. The microbial biomass exhibited a strong preference for ammonium over glycine and nitrate, regardless of invasion status. By comparison, native plants showed no nitrogen preferences and M. vimineum preferred inorganic nitrogen species. We interpret our findings as evidence that invasion by M. vimineum leads to changes in the partitioning of nitrogen above and belowground in forest understories, and to decreases in the microbial biomass, but it does not affect the outcome of plant–microbe–nitrogen interactions, possibly due to functional shifts in the microbial community as a result of invasion.     

Alteration of microbial community composition and changes in decomposition associated with an invasive intertidal macrophyte

Data demonstrating the effects of biological invaders on microbial communities and microbial processes are scarce, especially in marine environments. Research was conducted at Padilla Bay, Washington, to examine the effect that an invasive intertidal eelgrass, Zostera japonica Aschers & Graebn, has on rates of decomposition, microbial community composition, and the possible implications for associated ecosystem processes in this estuarine environment. A series of observational and experimental studies were conducted in beds of Z. japonica, beds of its native congener, Zostera marina, and mixed eelgrass beds. These studies assessed decomposition of invasive and native eelgrass, enumerated bacterial abundance, and examined sole source carbon usage (SSCU) by microbial assemblages. Z. japonica decomposed more rapidly than its native congener throughout the study period although rates of decomposition were variable. Microbial abundance did not differ among different vegetation compositions although differences in SSCU by microbial assemblages were detected among beds of invasive, native, and mixed eelgrass. These results indicate that this abundant invasive species can accelerate rates of decomposition and alter the associative decomposer community, which may lead to higher carbon and nutrient turnover within Padilla Bay.