Plant-soil feedbacks between invasive shrubs and native forest understory species lead to shifts in the abundance of mycorrhizal fungi

Aims

Non-native shrubs are important invaders of the Eastern Deciduous Forest, dramatically altering forest structure and functioning. Study of invasion mechanisms in this system has emphasized aboveground processes, and plant-soil feedbacks are relatively unexplored as a mechanism of shrub dominance. We tested whether plant-soil feedback in this habitat is affected by competition and whether arbuscular mycorrhizal fungi (AMF) are involved in plant-soil feedback.

Methods

We used a standard two-phase plant-soil feedback experiment run concurrently for each of three invasive shrub species, measuring feedback effects on AMF colonization, aboveground biomass, and the responses of native plant species in greenhouse mesocosms.

Results

Lonicera maackii and Ligustrum vulgare reduced AMF colonization of native roots, both with legacy effects (prior growth in soil) and direct effects (current growth in soil). Elaeagnus umbellata grown with natives left a legacy of increased AMF colonization of native communities.

Conclusions

Our results suggest that woody invasive species can alter the AMF associations of native plants even after the invasive is no longer present. Such consequences merit study with other native species and where environmental factors, such as light availability, might be expected to compound the effects of changes in AMF.     

Allelopathic potentials of exotic invasive and native trees over coexisting understory species: the soil as modulator

Ailanthus altissima and Robinia pseudoacacia are two aggressive exotic tree species invading riparian ecosystems in Central Spain. We explored their allelopathic potentials as a possible mechanism explaining their success in these ecosystems. Specifically, we aimed (1) to compare the phytotoxic effects of the exotic and native (Fraxinus angustifolia and Populus alba) trees on the fitness of several understory plants coexisting in riparian ecosystems, and (2) to assess the capacity of the riparian soil to modulate the phytotoxic effects. In laboratory bioassays, aqueous leaf litter extracts from the donor tree species at field-realistic concentrations were tested on different fitness indicators of 13 understory target species, using germination paper and soil as substrates in petri dishes. Using germination paper, we found species-specific effects between donor and target species, but the phytotoxicity of the exotic trees as a group was not greater than that of the natives. Nevertheless, the exotic R. pseudoacacia was the most effective donor species reducing the radicle growth of the target species. Over riparian soil substrate, the aqueous leaf litter extracts did not produce any phytotoxic effect on the target species, except in one case. Altogether, our results highlight the importance of using both a native control when assessing the phytotoxicity of nonnative plants and also the natural soil in the modulation of phytotoxic effects. Ignoring both factors in laboratory bioassays would have led to the overestimation of the phytotoxicity of the exotic species as a mechanism contributing to their invasion success.     

Elevated native terrestrial snail abundance and diversity in association with an invasive understory shrub,Berberis thunbergii,in a North American deciduous forest

Invasive terrestrial plants often substantially reshape environments, yet how such invasions affect terrestrial snail assemblages remains understudied. We investigated how snail assemblages in deciduous forest soils with dense Berberis thunbergii (Japanese barberry), an invasive shrub in eastern North America, differ from forest areas lacking the shrub. Leaf litter and soil samples were collected from forest patches with dense B. thunbergii understories and adjacent control areas within two exurban forest tracts in western Pennsylvania, U.S.A. Snails were identified to species and quantified by standard diversity metrics. Contrary to our expectations, snails were significantly more abundant and diverse in B. thunbergii-invaded areas. Despite differences in abundance, the snail community composition did not differ between invaded and control habitats. The terrestrial snail assemblage we observed, which was composed entirely of native species, appears to respond favorably to B. thunbergii invasion and therefore may not be negatively impacted by physicochemical changes to soils typically observed in association with the plant. Such findings could reflect the fact that B. thunbergii likely creates more favorable habitat for snails by creating cooler, more humid, and more alkaline soil environments. However, the snail assemblages we retrieved may consist mostly of species with high tolerance to environmental degradation due to a legacy of land use change and acid deposition in the region.     

Influence of woody invader control methods and seed availability on native and invasive species establishment in a Hawaiian forest

When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai’i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but short-lived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.     

Light, nutrients and the growth of herbaceous forest species

The herb layer of forests planted on former agricultural land often differs from that of old-growth forest. This study investigates if the expected increased nutrient availability in the shaded conditions of newly planted forests and the plasticity of the species to adjust their biomass allocation to different levels of light and nutrients help to explain these differences in the herb layers of the two forest types. In a greenhouse experiment biomass distribution and production of two species characteristic for the highly shaded forest floor, Circaea lutetiana and Mercurialis perennis, and two species more common in the forest-edge, Aegopodium podagraria and Impatiens parviflora were studied at different levels of light (2%, 8% and 66% of the full light level) and nutrients (30 and 300 kg N ha^–1 per year). The main factor affecting allocation and biomass production was light availability. Nutrient supply only had a significant effect at the higher light levels. Species were mainly plastic to changes in light and the two species from the forest floor showed to be more rigid in allocation pattern than the species from the forest-edge. So, although the species from the forest-edge were more plastic, they did not profit from the increased nutrient supply because the main factor affecting biomass distribution and production was light availability.     

Nesting bird species in sun coffee, pasture, and understory forest in southern Costa Rica

We investigated avian nest distribution and success in understoryforest, sun coffee plantations, and pasture in southern Costa Rica. Nestsearching occurred in plantations and forest in 1999 and 2000 and in pastures in2000. Nests were monitored until they failed or fledged young. Antbirds(Thamnophilidae) were the most common understory forest nesters and were notfound nesting in the plantations or pastures. Common nesting species in theplantations included Turdidae, Tyrannidae, Cardinalidae, and Thraupidae, many ofwhich are typical of forest edge/canopy or open, scrubby habitats. Two speciesassociated with forest interior, Henicorhina leucostictaand Buarremon brunneinucha, were found nesting in theplantations. Pastures supported similar types of nesting species as theplantations, with the exception of the forest-interior species. Daily mortalityrates (DMRs) for above-ground cup-nesting species in plantations and pastureswere similar to those for species nesting in forest at our site and a site inPanama. The results indicate that conversion from forest to pastures and suncoffee plantations diminishes nesting habitat for forest-interior species, whilenumerous forest edge/canopy species and open-country species are able to nest inthese agricultural land-cover types. As a group, species nesting in theplantations and pastures do not have unusually high nest mortality rates,although species-specific studies are lacking. Nesting species distributionsacross habitat types and DMRs at our study site may be influenced by the largeamount of forest in the landscape.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Thermal ecophysiology of a native and an invasive gecko species in a tropical dry forest of Mexico

For ectotherms, thermal physiology plays a fundamental role in the establishment and success of invasive species in novel areas and, ultimately, in their ecological interactions with native species. Invasive species are assumed to have a greater ability to exploit the thermal environment, higher acclimation capacities, a wider thermal tolerance range, and better relative performance under a range of thermal conditions. Here we compare the thermal ecophysiology of two species that occur in sympatry in a tropical dry forest of the Pacific coast of Mexico, the microendemic species Benedetti's Leaf-toed Gecko (Phyllodactylus benedettii) and the invasive Common House Gecko (Hemidactylus frenatus). We characterized their patterns of thermoregulation, thermoregulatory efficiency, thermal tolerances, and thermal sensitivity of locomotor performance. In addition, we included morphological variables and an index of body condition to evaluate their effects on the thermal sensitivity of locomotor performance in these species. Although the two species had similar selected temperatures and thermal tolerances, they contrasted in their thermoregulatory strategies and thermal sensitivity of locomotor performance. Hemidactylus frenatus had a higher performance than the native species, P. benedettii, which would represent an ecological advantage for the former species. Nevertheless, we suggest that given the spatial and temporal limitations in habitat use of the two species, the probability of agonistic interactions between them is reduced. We recommend exploring additional biotic attributes, such as competition, behavior and niche overlap in order assess the role of alternative factors favoring the success of invasive species.     

No universal scale-dependent impacts of invasive species on native plant species richness

A growing number of studies seeking generalizations about the impact of plant invasions compare heavily invaded sites to uninvaded sites. But does this approach warrant any generalizations? Using two large datasets from forests, grasslands and desert ecosystems across the conterminous United States, we show that (i) a continuum of invasion impacts exists in many biomes and (ii) many possible species–area relationships may emerge reflecting a wide range of patterns of co-occurrence of native and alien plant species. Our results contradict a smaller recent study by Powell et al. 2013 (Science 339, 316–318. (doi:10.1126/science.1226817)), who compared heavily invaded and uninvaded sites in three biomes and concluded that plant communities invaded by non-native plant species generally have lower local richness (intercepts of log species richness–log area regression lines) but steeper species accumulation with increasing area (slopes of the regression lines) than do uninvaded communities. We conclude that the impacts of plant invasions on plant species richness are not universal.     

Bird species diversity in forest understory: Analysis of mist-net samples

Summary Mist-netted samples of forest understory birds in neotropical (Costa Rica) and temperate (Illinois) forests (during spring migration) supported certain earlier observations: the neotropical understory sample contained more species, more rare species, and more species that use resources not sufficiently available in temperate forest.During spring migration in Illinois, the number of bird species in a mist-netted sample of the community approaches (albeit temporarily) that of neotropical forests and is sometimes even equivalent. But the constitution of these samples seems to be quite different in different regions. Costa Rican samples showed greater within-habitat variability and individual recapture distances were shorter, implying a greater local patchiness of bird distributions than in Illinois in spring.     

Predicting the potential distribution and forest impact of the invasive species Cydalima perspectalis in Europe

Invasive species have considerably increased in recent decades due to direct and indirect effects of ever‐increasing international trade rates and new climate conditions derived from global change. We need to better understand how the dynamics of early species invasions develop and how these result in impacts on the invaded ecosystems. Here we studied the distribution and severe defoliation processes of the box tree moth (Cydalima perspectalis W.), a tree defoliator insect native to Asia and invasive in Europe since 2007, through the combination of species distribution models based on climate and landscape composition information. The results showed that the combination of data from the native and the invaded areas was the most effective methodology for the appropriate invasive species modeling. The species was not influenced by overall landscape factors, but only by the presence of its host plant, dispersal capacity, and climate suitability. Such climate suitability was described by low precipitation seasonality and minimum annual temperatures around 0°C, defining a continentality effect throughout the territory. We emphasize the need of studying distribution and severe defoliation processes separately because we identified that climate suitability was slightly involved in limiting species spread processes but strongly constrained ecosystem impact in terms of defoliation before the species reaches equilibrium with the new environment. New studies on habitat recovery after disturbance, ecological consequences of such impact, and community dynamics in a context of climate change are required for a better understanding of this invasive species.     

The effects of nitrogen addition on the growth of two exotic and two native forest understory plants

Many studies have linked the spread of exotic, invasive species to high nitrogen supply, but most of this work does not distinguish between various inorganic forms and different concentrations of available nitrogen. Previous research has suggested that exotic, invasive species common in eastern deciduous forests may preferentially utilize nitrate in contrast to native species that preferentially make use of ammonium. We tested this hypothesis by comparing the growth response of two common forest invaders, Berberis thunbergii and Microstegium vimineum, and two co-occurring native species (Vaccinium pallidum and Hamamelis virginiana) to different forms of nitrogen under varying concentrations in a greenhouse experiment. Two forms of nitrogen (nitrate or ammonium) were added at different concentrations (22, 106, and 212 mg N l⁻¹) to all species. Growth response variables included survival, stem length, stem diameter, above and belowground biomass, and estimated seed production (Microstegium only). Unexpectedly the exotic species did not respond preferentially to nitrate addition. Microstegium responded most strongly to both nitrogen forms depending on the response variable. Berberis and Hamamelis surprisingly reacted similarly to nitrogen additions. As expected, Vaccinium fared poorly under most treatment conditions, but did show some growth in NH₄ ⁺ treatments. Our findings suggest species response to nitrogen addition is complex, and that exotic species do not all respond similarly to nitrogen inputs. In this study, the response of exotic and native species to available nutrients does not provide a general mechanism of invasion success.     

蕨类植物的化感作用及其对生物多样性的影响

植物化感作用是植物通过向环境中释放化学物质从而对同种植株和繁殖体或与其他植物之间产生的直接或间接、有益或有害的作用,它影响植物分布、群落形成与演化、作物间作等,与生物多样性保护以及农林和园艺生产实践关系密切,在国际上受到越来越多的关注。但国内这方面的研究起步相对较晚,研究范围有限,对蕨类植物化感作用的报道更少。本文系统介绍了蕨类植物化感作用的研究进展,包括蕨类植物种内的化感作用(即自毒效应)、常见的蕨类植物种间的化感作用(即孢子体对配子体的化感作用和配子体对配子体的化感作用)及蕨类植物对种子植物的化感作用(蕨类植物可通过化感作用与种子植物争夺更多的资源和生长空间)。还介绍了种子植物对蕨类植物的化感作用(主要表现为抑制作用)以及蕨类植物化感作用与动物侵食、微生物侵染的关系,研究发现昆虫侵食能增强或减弱蕨类植物的化感作用,微生物的活动可能增强某些蕨类植物的化感作用。本文从上述不同角度说明蕨类植物化感作用对生物多样性的影响,希望有助于促进我国学者对该领域的深入研究。     

Growth, biomass allocation and photosynthesis of invasive and native Hawaiian rainforest species

Growth, biomass allocation, and photosynthetic characteristics of seedlings of five invasive non-indigenous and four native species grown under different light regimes were studied to help explain the success of invasive species in Hawaiian rainforests. Plants were grown under three greenhouse light levels representative of those found in the center and edge of gaps and in the understory of Hawaiian rainforests, and under an additional treatment with unaltered shade. Relative growth rates (RGRs) of invasive species grown in sun and partial shade were significantly higher than those for native species, averaging 0.25 and 0.17 g g⁻¹ week⁻¹, respectively, while native species averaged only 0.09 and 0.06 g g⁻¹ week⁻¹, respectively. The RGR of invasive species under the shade treatment was 40% higher than that of native species. Leaf area ratios (LARs) of sun and partial-shade-grown invasive and native species were similar but the LAR of invasive species in the shade was, on average, 20% higher than that of native species. There were no differences between invasive and native species in biomass allocation to shoots and roots, or in leaf mass per area across light environments. Light-saturated photosynthetic rates (Pmax) were higher for invasive species than for native species in all light treatments. Pmax of invasive species grown in the sun treatment, for example, ranged from 5.5 to 11.9 μmol m⁻² s⁻¹ as compared with 3.0−4.5 μmol m⁻² s⁻¹ for native species grown under similar light conditions. The slope of the linear relationship between Pmax and dark respiration was steeper for invasive than for native species, indicating that invasive species assimilate more CO₂ at a lower respiratory cost than native species. These results suggest that the invasive species may have higher growth rates than the native species as a consequence of higher photosynthetic capacities under sun and partial shade, lower dark respiration under all light treatments, and higher LARs when growing under shade conditions. Overall, invasive species appear to be better suited than native species to capturing and utilizing light resources, particularly in high-light environments such as those characterized by relatively high levels of disturbance. Received: 30 December 1997 / Accepted: 1 September 1998     

Soil modification by invasive plants: effects on native and invasive species of mixed-grass prairies

Invasive plants are capable of modifying attributes of soil to facilitate further invasion by conspecifics and other invasive species. We assessed this capability in three important plant invaders of grasslands in the Great Plains region of North America: leafy spurge (Euphorbia esula), smooth brome (Bromus inermis) and crested wheatgrass (Agropyron cristatum). In a glasshouse, these three invasives or a group of native species were grown separately through three cycles of growth and soil conditioning in both steam-pasteurized and non-pasteurized soils, after which we assessed seedling growth in these soils. Two of the three invasive species, Bromus and Agropyron, exhibited significant self-facilitation via soil modification. Bromus and Agropyron also had significant facilitative effects on other invasives via soil modification, while Euphorbia had significant antagonistic effects on the other invasives. Both Agropyron and Euphorbia consistently suppressed growth of two of three native forbs, while three native grasses were generally less affected. Almost all intra- and interspecific effects of invasive soil conditioning were dependent upon presence of soil biota from field sites where these species were successful invaders. Overall, these results suggest that that invasive modification of soil microbiota can facilitate plant invasion directly or via ‘cross-facilitation’ of other invasive species, and moreover has potential to impede restoration of native communities after removal of an invasive species. However, certain native species that are relatively insensitive to altered soil biota (as we observed in the case of the forb Linum lewisii and the native grasses), may be valuable as ‘nurse’species in restoration efforts.     

Light use efficiency of California redwood forest understory plants along a moisture gradient

We investigated photosynthesis of five plant species growing in the understory at three sites (1,170-, 1,600- and 2,100-mm annual moisture inputs), along the geographical range of coastal California redwood forest, to determine whether greater inputs of rain and fog at northern sites enhance photosynthetic utilization of fluctuating light. Measurements of understory light environment and gas exchange were carried out to determine steady state and dynamic photosynthetic responses to light. Leaf area index ranged from 4.84 at the 2,100-mm site to 5.98 at the 1,170-mm site. Maximum rates of net photosynthesis and stomatal conductance (g) did not vary appreciably within species across sites. Photosynthetic induction after a change from low to high light was significantly greater in plants growing in lower light conditions regardless of site. Photosynthetic induction also increased with the rate of g in diffuse light, prior to the increase to saturating light levels. Post-illumination CO₂ assimilation was the largest factor contributing to variation in C gain during simulated lightflecks. The duration of post-illumination photosynthetic activity, total CO₂ assimilation per light received, and light use efficiency during simulated lightflecks increased significantly with moisture inputs in four out of five species. Increasing leaf N concentration with increasing moisture inputs in three out of five species, coupled with changes in leaf N isotopic composition with the onset of the summer fog season suggest that natural N deposition increases with rain and fog inputs and contributes to greater utilization of fluctuating light availability in coastal California redwood forests.