An invasive macrophyte alters sediment chemistry due to suppression of a native isoetid

The submersed macrophyte Utricularia inflata (inflated bladderwort) is a recent invader of Adirondack Mountain lakes (NY, USA). A 15-week greenhouse experiment and a 7-week field experiment were conducted to test the hypothesis that this rootless species fundamentally changes sediment chemistry through its suppression of the native short-statured species, Eriocaulon aquaticum. E. aquaticum has an extensive root system that releases oxygen into the sediment. In greenhouse conditions, E. aquaticum raised the porewater redox potential of otherwise bare sediment from 25 to 324 mV, lowered the sediment porewater pH from 5.7 to 4.6, and depleted the dissolved inorganic carbon and ammonium concentrations in the sediment porewater by 68.4 and 96.0%, respectively (P<0.001 for all four parameters). A cover of U. inflata over E. aquaticum, however, greatly reduced the latter’s effect on redox potential (P<0.001), dissolved solutes (P<0.001), and pH (P<0.05). E. aquaticum biomass increased during the greenhouse experiment in the absence of U. inflata, but decreased in its presence (P<0.001). Redox and growth rate results from the field experiment paralleled those from the greenhouse experiment. Our data suggest that U. inflata may change nutrient cycling in Adirondack lake ecosystems by reducing the growth of native isoetid macrophytes, such as E. aquaticum, and consequently altering key features of sediment chemistry.     

Exposure provides refuge from a rootless invasive macrophyte

Free-floating Utricularia inflata Walt. has the potential to alter native isoetid communities as it expands its range from the Atlantic coastal plain into northeastern New York lakes. This field study assessed (1) in situ growth potential for U. inflata at varying depths (1.0-3.0 m), (2) displacement of U. inflata from sites with different exposures to wind and water movement, and (3) U. inflata's relative abundance over a depth range (0.5 m, 1.0 m, 1.5 m, 2.0 m, and 2.5 m) at each of nine Adirondack Mountain lake sites varying in exposure. Plants grew well in shallow water (RGR: 0.014-0.039 day⁻¹) at depths from 1.0 m to 2.5 m, but lost mass at 3.0 m. Significantly fewer U. inflata plants remained at sites and depths with greater exposure to wave action, as well as those with greater water current. Vegetation sampling confirmed greater relative frequency of U. inflata at sheltered sites and along deeper contours. Despite its potential to grow well in shallow water, water movement can prevent the accumulation of U. inflata there, and thus provide a refuge for native species able to withstand wave exposure.     

Transpiration by an emergent macrophyte: source of water and implications for nutrient supply

The contribution of sediment interstitial water and the water column to the transpiration stream of Myriophyllum aquaticum (Vellozo) Verdcourt was determined to estimate the significance of mass flow in supply of sediment nutrients for plant growth. Sediment interstitial water accounted for about 2% of the water transpired over a 37 day period. Because of the small volume of water that originated in the sediment we concluded that mass flow did not significantly enhance nutrient supply to the roots of M. aquaticum. Relative growth rate (RGR) of adventitious, water roots was greater than whole plant RGR, and RGR of sediment roots was not significantly different from zero, indicating a shift in the biomass allocation after emergence of the apical meristem into the air. Water use, measured by the transpiration coefficient, averaged 260 ml H₂O mg DW^-1, which is similar to C-4 terrestrial plants. M. aquaticum has leaf characteristics commonly associated with xerophytic habitats. These characteristics may be necessary if a high transpiration rate and a mechanical requirement for high cell turgor pressure, required by a reliance upon hydrostatic pressure for support of the aerial stems, are mutually exclusive because of morphological constraints on hydraulic conductivity.     

Apparent facilitation of an invasive mealybug by an invasive ant

Summary In the southeast United States, the invasive ant Solenopsis invicta is known to derive important carbohydrate (honeydew) resources from mealybugs utilizing grasses. Most important appears to be an invasive mealybug, Antonina graminis. We studied whether this mealybug and a similar native species also benefit from association with S. invicta. We found that mealybug occurrence increases significantly with increasing proximity to S. invicta mounds, suggesting that mealybugs benefit as well. Mutual benefits derived by S. invicta and A. graminis are consistent with a hypothesis proposing that associations among invasive species can be important in their success at introduced locations.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

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.     

Habitat modification inhibits conspecific seedling recruitment in populations of an invasive ecosystem engineer

Invasive species that strongly modify their physical habitat are a particular management concern. Theoretical models predict that habitat modification could speed spread rates or allow invasion of sites that would otherwise resist invasion. There are few empirical tests of this hypothesis, however. We tested whether habitat modification by invading Spartina alterniflora populations facilitates conspecific seedling recruitment and spatial spread in Willapa Bay, WA, USA. Established S. alterniflora individuals strongly modified their local physical environment. Hydrologic flow, porewater salinity, and light availability were decreased while sediment NH₄ ⁺ increased with increasing S. alterniflora stem density. The S. alterniflora seed bank was greater and spring seedlings were denser within meadows of S. alterniflora than on unvegetated tideflats. However, almost all seedling recruitment after 1 year occurred on tideflats or on meadow edge plots where the above ground S. alterniflora biomass had been removed. Instead of facilitating invasive spread, ecosystem engineering in this system appears to create conditions that inhibit local seedling recruitment. These results suggest that the influence of ecosystem engineering on invasive spread is highly contingent on the relative spatial scales of habitat modification, environmental heterogeneity, and propagule availability. Control activities could change these spatial relationships, however, inadvertently promoting invasive recruitment.     

遮阳对白花前胡叶片光合特性的影响

为规范白花前胡(Peucedanum praerupterum)种植和提高产量,对不同光强下白花前胡的光合特性进行了研究.结果表明,与遮阳40％和60％相比,对照(CK)和遮阳20％的白花前胡叶片具有更高的最大电子传递速率[分别为247.129和266.866μmol/(m2·s)]和最大净光合速率[分别为25.621...     

A Panax-centric view of invasive species

Plant-centric sampling provides a novel approach to quantifying the potential impact of invasive species on native plant species. The aim of this study was to determine the level of exposure of individuals and populations of Panax quinquefolius to invasive plant species using this approach in thirty natural ginseng populations. A high level of invasion was found with 63–70% of ginseng populations containing at least one invasive species. Approximately one-third of all individuals were found in close proximity to invasive plants. The most prevalent invasive species were Rosa multiflora and Berberis thunbergii. The exposure to invasives of plants in different size classes varied among populations. Invasive species presence increased with greater ginseng population sizes and presence of harvest. The abundance of invasives plants within forest interiors near this valuable medicinal herb suggests that the economic and ecological costs of competitive interactions with native species could be high.     

Phosphorus fluctuation in water and deposition into sediment within an emergent macrophyte stand

Seasonal fluctuation of phosphorus in water, and total phosphorus and different inorganic P fractions (extracted by NH₄Cl, NH₄F, NaOH and H₂SO₄) and organic P fraction (residual P) in surface sediment, were measured in the littoral of oligotrophic Lake Pääjärvi (southern Finland). After the emergence of Equisetum fluviatile L. shoots in mid June, water exchange between the littoral and pelagial area diminished and phosphorus concentrations in water and in surface sediment increased in the inner and mid littoral zones. Phosphorus pool in flocculent, easily resuspensible sediment composed on average 62% of inorganic phosphorus and 38% of organic phosphorus. 63% of the inorganic phosphorus, on the average, was extracted from apatite-P fraction 29% from iron-P fraction 7% from aluminum-P fraction and less than 1% from loosely-bound P fraction. During the growing season, net accumulation of sediment and phosphorus was measured only in the inner littoral zone whereas the outer littoral zones acted rather as a source of phosphorus for the pelagic area. The results of this study indicated that nutrient dynamics in littoral environment was through changes in flow environment greatly governed by the macrophytes.     

The effects of nitrate loading on the invasive macrophyte Hydrilla verticillata and two common, native macrophytes in Florida

Rising nitrate concentrations in the water column and the spread of invasive, non-native macrophytes are two major threats to Florida's oligotrophic, freshwater ecosystems. We used a replicated mesocosm experiment to test the effects of elevated nitrate concentrations in the water on the growth of the invasive macrophyte Hydrilla verticillata and two common, native submerged macrophytes Vallisneria americana and Sagittaria kurziana. Results from this study indicate that nitrate concentrations of 1.0 mg L⁻¹ NO₃-N in the water increased the final dry-weight biomass of H. verticillata by 2.75 times, while having no statistical effect on the growth of the two native species. Additionally, H. verticillata grew at a faster rate than the two native species in the low nitrate treatments accounting for 82% of the total biomass, indicating that it may have the capacity to invade relatively pristine communities. In waters where nitrate concentrations continue to rise, the cost of control efforts for H. verticillata may substantially increase in the future.     

Geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata

We investigated geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata, to test the hypothesis that escape from herbivory in invasive species permits enhanced growth and lower production of defensive chemicals. We quantified herbivore damage, concentrations of sinigrin, and growth and reproduction inside and outside herbivore exclusion treatments, in field populations in the native and invasive ranges. As predicted, unmanipulated plants in the native range (Hungary, Europe) experienced greater herbivore damage than plants in the introduced range (Massachusetts and Connecticut, USA), providing evidence for enemy release, particularly in the first year of growth. Nevertheless, European populations had consistently larger individuals than US populations (rosettes were, for example, eightfold larger) and also had greater reproductive output, but US plants produced larger seeds at a given plant height. Moreover, flowering plants showed significant differences in concentrations of sinigrin in the invasive versus native range, although the direction of the difference was variable, suggesting the influence of environmental effects. Overall, we observed less herbivory, but not increased growth or decreased defense in the invasive range. Geographical differences in performance and leaf chemistry appear to be due to variation in the environment, which could have masked evolved differences in allocation.     

遮阴对四季桂生理生态特性的影响

为了解遮阴对四季桂(Osmanthus fragrans)生长的影响,研究了成都地区夏季遮阴处理下四季桂扦插苗生理生态特性的变化。结果表明,随遮光率增加,比叶重下降;叶绿素b(Chl b)含量增加,Chl a/b降低。遮阴下叶片可溶性糖、可溶性蛋白含量降低,以80%遮光率下降的最显著;随遮光率上升,叶片丙二醛(MDA)含量下降,超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)活性降低,以55%、80%遮光率的MDA含量下降显著。遮阴处理下,叶片光饱和点(LSP)、光补偿点(LCP)、最大净光合速率(Pmax)、暗呼吸速率(Rd)均下降,而表观量子速率(AQE)增大;瞬时光能利用率(LUE)升高,净光合速率(Pn)、气孔导度(Gs)、气孔限制值(Ls)均降低,胞间CO_2浓度(Ci)升高。四季桂幼苗主要通过提高Pn和抗氧化酶活性来适应强光环境;降低SLW、LSP、LCP和Rd,提高Chl和Ci含量,增大AQE和LUE提高弱光利用能力,中度遮阴(25%、55%遮光率)有利于幼苗在夏季的生长。     

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.     

Lethal control of an apex predator has unintended cascading effects on forest mammal assemblages

Disruption to species-interaction networks caused by irruptions of herbivores and mesopredators following extirpation of apex predators is a global driver of ecosystem reorganization and biodiversity loss. Most studies of apex predators'' ecological roles focus on effects arising from their interactions with herbivores or mesopredators in isolation, but rarely consider how the effects of herbivores and mesopredators interact. Here, we provide evidence that multiple cascade pathways induced by lethal control of an apex predator, the dingo, drive unintended shifts in forest ecosystem structure. We compared mammal assemblages and understorey structure at seven sites in southern Australia. Each site comprised an area where dingoes were poisoned and an area without control. The effects of dingo control on mammals scaled with body size. Activity of herbivorous macropods, arboreal mammals and a mesopredator, the red fox, were greater, but understorey vegetation sparser and abundances of small mammals lower, where dingoes were controlled. Structural equation modelling suggested that both predation by foxes and depletion of understorey vegetation by macropods were related to small mammal decline at poisoned sites. Our study suggests that apex predators’ suppressive effects on herbivores and mesopredators occur simultaneously and should be considered in tandem in order to appreciate the extent of apex predators’ indirect effects.     

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.     

Ladybird population dynamics in potato: comparison of native species with an invasive species, Harmonia axyridis

Following the detection of the harlequin ladybird, Harmonia axyridis, in 2003 in potato crops in Belgium, a study was carried out between 2004 and 2006 on the phenology of this species compared to native species in potato. The results confirmed the success of H. axyridis, with high population levels in 2004 and 2005. In 2006, aphid populations were very low and no H. axyridis larvae were sampled in potato, but the indigenous species Coccinella septempunctata and Propylea quatuordecimpunctata were detected. A species by species comparison of the date of first larvae detection, the larvae population peak, and the difference between this peak and the aphid population peak was performed. Results showed a clear correlation between C. septempunctata and P. quatuordecimpunctata and potato aphids, with a delay of 3.5 and 6.5 days between the aphid and ladybird population peaks for the two native species. H. axyridis arrived 7–8 days after the two indigenous species and the larval peak population occurred 15.8 days after the aphid population peak. This meant that H. axyridis had to complete its larval development with very low aphid populations or even with no aphids at all. The reason for its late arrival and the possible food resources used by H. axyridis larvae are discussed.     

Ex situ evaluation of impacts of invasive mosquitofish on the imperiled Barrens topminnow

Central to the protection of native species is an understanding of impacts of actual or potential invasive species and also the mechanisms through which those impacts are mediated. The introduction and spread of western mosquitofish, Gambusia affinis, into spring systems of the Barrens Plateau region of middle Tennessee is a concern for native species such as the Barrens topminnow, Fundulus julisia. We investigated whether mosquitofish might act as predators on early life stages of topminnows as well as affect the physiological well being of adults through aggressive interactions. A short-term, 24-h laboratory study with mosquitofish and topminnows demonstrated the vulnerability of young topminnow life stages to large mosquitofish predation and aggression. Survival of topminnow young, <16 mm total length (TL), was 0% and was attributed to predation by mosquitofish. Survival of juveniles, 20–30 mm TL, was 25%; juveniles mostly succumbed (post 24-h) to injuries inflicted by large mosquitofish. Adult topminnow survival was 100% but adults faced injury risk, primarily during the initial stages of their interaction with large mosquitofish. A long-term, 60-day laboratory study with syntopic and allotopic populations of adult topminnows and mosquitofish failed to detect any negative impacts on topminnows due to coexistence. Survival, growth, and fecundity of adult topminnows syntopic with mosquitofish were not different from the allotopic population, although injury risk in the form of fin damage was greater syntopically. Thus, predation and aggression towards young topminnows may be the primary mechanisms by which western mosquitofish jeopardize the persistence of native Barrens topminnows in the wild. Our results reemphasize the danger to native aquatic biodiversity of unregulated introductions of Gambusia species.     

Solitary invasive orchid bee outperforms co-occurring native bees to promote fruit set of an invasive Solanum

Our understanding of the effects of introduced invasive pollinators on plants has been exclusively drawn from studies on introduced social bees. One might expect, however, that the impacts of introduced solitary bees, with much lower population densities and fewer foragers, would be small. Yet little is known about the potential effects of naturalized solitary bees on the environment. We took advantage of the recent naturalization of an orchid bee, Euglossa viridissima, in southern Florida to study the effects of this solitary bee on reproduction of Solanum torvum, an invasive shrub. Flowers of S. torvum require specialized buzz pollination. Through timed floral visitor watches and two pollination treatments (control and pollen supplementation) at three forest edge and three open area sites, we found that the fruit set of S. torvum was pollen limited at the open sites where the native bees dominate, but was not pollen limited at the forest sites where the invasive orchid bees dominate. The orchid bee’s pollination efficiency was nearly double that of the native halictid bees, and was also slightly higher than that of the native carpenter bee. Experiments using small and large mesh cages (to deny or allow E. viridissima access, respectively) at one forest site indicated that when the orchid bee was excluded, the flowers set one-quarter as many fruit as when the bee was allowed access. The orchid bee was the most important pollinator of the weed at the forest sites, which could pose additional challenges to the management of this weed in the fragmented, endangered tropical hardwood forests in the region. This specialized invasive mutualism may promote populations of both the orchid bee and this noxious weed. Invasive solitary bees, particularly species that are specialized pollinators, appear to have more importance than has previously been recognized. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

遮阴对地枫皮幼苗生长和生物量分配的影响

为了解光照强度对地枫皮(Illicium difengpi)幼苗生长和生物量分配的影响,对不同等级幼苗在不同遮阴处理下的形态和生物量变化进行了研究。结果表明,遮阴处理下幼苗的株高、冠宽和平均单叶面积均显著高于对照,同时比叶面积均随光强增大而显著降低。除I级苗50%遮阴处理外,全光处理下各等级幼苗的根长均显著高于遮阴处理。随着光照强度的增加,各等级幼苗的根生物量比与根冠比显著增加,叶生物量比显著减少;不同等级幼苗对强光和弱光环境均表现出较强的适应性,但以50%遮阴处理的总生物量最大,全光和85%遮阴处理均较小。遮阴处理的幼苗个体大小与其初始大小呈正相关,其中II级苗与I级苗差别较小,III级苗生长较差。在人工育苗条件下I级苗数量不到10%,较差的种苗质量叠加喀斯特山顶恶劣环境,可能是导致野外幼苗更新限制的重要原因。因此在野外回归和人工栽培过程中宜选用I、II级种苗,光强控制在透光率50%为宜。