Growth of Phragmites australis and Phalaris arundinacea in constructed wetlands for wastewater treatment in the Czech Republic

Common reed (Phragmites australis) and reed canarygrass (Phalaris arundinacea) are two most commonly used plant species in constructed wetlands for wastewater treatment in the Czech Republic. Growth characteristics of both plants (biomass, stem count, and length) have been measured in 13 horizontal sub-surface flow constructed wetlands since 1992. The results revealed that while Phalaris usually reaches its maximum biomass as early as during the second growing season, Phragmites usually reaches its maximum only after three to four growing seasons. The maximum biomass of both species varies widely among systems and the highest measured values (5070 g m⁻² for Phragmites and 1900 g m⁻² for Phalaris) are similar to those found in eutrophic natural stands. The shoot count of Phragmites decreases after the second growing season while length and weight of individual shoots increases over time due to self-thinning process. Number of Phalaris shoots is the highest during the second season and then the shoot count remains about the same. Also the shoot length remains steady over years of constructed wetland operation.     

Understanding invasion as a process: the case of <Emphasis Type="Italic">Phalaris arundinacea</Emphasis> in wet prairies

Invasive plants that most threaten biodiversity are those that rapidly form a monospecific stand, like the clonal grass, Phalaris arundinacea. Understanding complex and potentially interacting factors that are common in urban and agricultural landscapes and underlie rapid invasions requires an experimental, factorial approach. We tested the effects of flooding and nutrient and sediment additions (3 × 3 × 3 = 27 treatments, plus a control with no additions) on invasion of Phalaris into mesocosms containing wet prairie vegetation. We discovered a three-step invasion and degradation process: (1) initially, resident native species declined with prolonged flooding and sediment additions, and (2) prolonged flooding, sedimentation, and nutrients accelerated Phalaris aboveground growth; biomass rose to 430 times that of the control within just two growing seasons. The dramatic expansion of Phalaris in the second year resulted in the formation of monospecific stands in over one-third of the treatments, as (3) native species continued their decline in year 2. Disturbances acted alone and in combination to make the resident wetland community more invasible and Phalaris more aggressive, leading to monospecific stands. Yet, Phalaris did not always “win”: under the least disturbed conditions, the resident plant canopy remained dense and vigorous and Phalaris remained small. When anthropogenic disturbances coincide with increases in the gross supply of resources, more tolerant, fast-growing, and morphologically plastic plants like Phalaris can invade very rapidly. The fluctuating resource hypothesis should thus be refined to consider the role of interacting disturbances in facilitating invasions.     

Set‐backs in Replacing Phalaris arundinacea Monotypes with Sedge Meadow Vegetation

Reed canary grass (Phalaris arundinacea) invades wetlands, forms monotypes, and resists control efforts, suggesting that strong feedbacks sustain its dominance, as in the alternative states model. In nine field experiments, we tested the hypothesis that applying a graminicide (sethoxydim) for three years would progressively reduce Phalaris abundance, and that seeding sedge meadow species (except grasses) would reestablish native plant dominance. The graminicide prevented Phalaris from flowering, reduced its height by 50% and reduced its cover, often to less than 40%. However, only two of the nine sites showed progressive declines over the three‐year experiment. The first setback was that Phalaris recovered annually in nearly all treatment plots. A second setback was that seeding did not reestablish sedge meadow. In five sites, unseeded plots had similar numbers of native species as those seeded with either forbs, forbs and graminoids, or graminoids. In four formerly agricultural sites, however, non‐native weeds increased in species richness and cover (a third setback). In only one site did the graminicide's effect on Phalaris allow native species to increase in number and cover. But short‐term gains were not long‐lasting. In year four, three sites that developed high native‐species cover were again strongly dominated by Phalaris (a fourth setback). The feedbacks that sustain this invader include resistance to the graminicide aboveground and rapid and robust regrowth from rhizomes and seeds belowground. The weak effect of this graminicide was a surprise; hence, we recommend stronger management actions to control Phalaris.     

本地和外来草本物种对水分条件时间异质性的可塑响应

极端气候导致的干旱和水淹事件频发，影响了外来植物和本地植物的生长。为了解外来种和本地种植物对干旱和水淹事件发生顺序的响应，探讨草本植物适应水分时间异质性的策略，该文以美国蒙大拿州西部4种本地植物和4种外来植物为研究对象，将所有植物分别进行持续湿润(对照，CK)、水淹-干旱(I-D)和干旱-水淹(D-I)处理，并观测一系列形态和生物量特征的变化。结果表明：(1)与CK相比，D-I和I-D处理均显著降低了外来种的总生物量(P<0.05)。(2)D-I显著降低了本地种早期总生物量、后期地下生物量和根冠比，但显著提高了其后期的相对生长(P<0.05)。(3)D-I处理显著降低了所有植物的地下-地上生物量关系的异速指数，外来种异速指数显著高于本地种(P<0.05)。综上认为，极端事件(水淹和干旱)的发生顺序能改变外来植物和本地植物的生物量分配，早期干旱比后期干旱更容易减少植物生物量的积累，但能促进本地种后期的生长；本地种在环境胁迫下不被降低的总生物量表现说明维持表型稳定的能力较强；D-I处理下本地种和外来种地上和地下生物量关系的分配方式不同。     

3种本地植物与入侵植物紫茎泽兰的竞争

为筛选替代控制紫茎泽兰的本地植物及探讨提高替代控制效率的方法,通过盆栽实验并利用相对产量(RY)和竞争攻击力系数(A)衡量了3种本地植物与紫茎泽兰的竞争关系,同时评估了活性炭(AC)、杀真菌剂(FC)和二者联合(AC+FC)对它们竞争的影响。结果表明：(1)不添加任何物质条件下：紫茎泽兰与南酸枣混种时,紫茎泽兰的生物量明显高于单种(P<0.05),其RY和A分别显著大于1和0(P<0.05);紫茎泽兰分别与假地豆和狗尾草混种时,紫茎泽兰的株高和生物量均明显低于单种(P<0.05),其RY和A均分别显著小于1和0(P<0.05)。说明紫茎泽兰的竞争力强于南酸枣而弱于假地豆和狗尾草,假地豆和狗尾草可以在一定区域作为替代控制紫茎泽兰的潜在目标植物。(2)与不添加任何物质相比,紫茎泽兰与南酸枣混种时,AC、FC及AC+FC处理增加了紫茎泽兰的根冠比,降低了其地上生物量比,也降低了南酸枣的生物量(P<0.05);紫茎泽兰与假地豆混种时,AC和AC+FC处理增加了假地豆的株高和生物量,FC处理增加了紫茎泽兰的株高(P<0.05);紫茎泽兰与狗尾草混种时,AC和A...     

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.     

Does morphological plasticity of the <Emphasis Type="Italic">Phalaris arundinacea</Emphasis> canopy increase invasiveness?

Morphological plasticity could facilitate invasions of wetland plants into areas that experience increased durations of flooding and eutrophication. We explored canopy plasticity of Phalaris arundinacea, an aggressive invader of wetlands, as it differentially invaded wet prairie mesocosms under 3 flooding durations and 3 levels of nutrient addition. Phalaris grew as a sward with intermittent and early-season flooding but shifted to tussocks under constant flooding. These two growth forms differed by >20% in several canopy ratios. Clones that formed tussocks produced 45% more shoots per unit biomass (P = 0.007) and a 25% higher ratio of total shoot length to biomass (P = 0.04). Lighter-weight shoots supported 33% fewer leaves and, consequently, had 35% less leaf area per shoot height (P < 0.002). Tussocks developed a continuous mat of adventitious roots, with root mats reaching 20.9 ± 0.6 cm in diameter and 4.7 ± 0.3 cm in height over two growing seasons. While forming tussocks, Phalaris tolerated longer durations of flooding and more than doubled its aboveground biomass. Invasions occurred rapidly, with Phalaris exceeding 75% canopy cover and accounting for 66% of the total aboveground biomass under constant flooding. Early-season flooding increased the lateral spread of individual shoots. High nutrient addition produced shoots that were 27% taller and 50% heavier (P < 0.02), with 81% more leaf area (P < 0.0003) than shoots that received no nutrients. Consequently, under early-season flooding with high nutrient additions, Phalaris was primed to invade, nearly doubling its proportion of the total aboveground biomass and exceeding 50% canopy cover during year two.     

Exotic Spartina alterniflora provides compatible habitats for native estuarine crab Sesarma dehaani in the Yangtze River estuary

Although the impact of plant invasions on benthic communities, especially burrowing crabs, has received increasing attention, the results from past studies are mixed. The exotic plant Spartina alterniflora has become the most abundant species in the salt marshes of the Yangtze River estuary since it was first found just over a decade ago, but its effects on crabs in the salt marshes is largely unknown. To examine whether the invasions of this exotic plant affected native crabs, we compared the biomass and abundance of the dominant burrowing crab Sesarma dehaani in an exotic Spartina marsh, native Phragmites australis marsh and mudflats of the Yangtze River estuary, China. To explain the differences of S. dehaani populations between different habitats, feeding preference of S. dehaani for Spartina and Phragmites was investigated. Results showed crab abundance and biomass in the Spartina marsh were significantly greater than those in the Phragmites marsh and mudflats. Soil water content and plant community characteristics in the Spartina marsh also significantly differed in the Phragmites marsh and mudflats. Moreover, the feeding preference experiment showed that crabs consumed Spartina more than twice as much as Phragmites. In summary, this study showed that Spartina provided compatible habitats for native crab S. dehaani through offering suitable food source and moderate environmental conditions.     

Response of the invader Cortaderia selloana and two coexisting natives to competition and water stress

Alien species’ resistance and adjustment to water stress and plant competition might largely determine the success of invasions in Mediterranean ecosystems because water availability is often limiting biomass production. Two outdoor pot experiments were conducted to test the hypotheses that at the recruitment stage the invader perennial tussock grass Cortaderia selloana is a superior competitor, and that it is more resistant to water stress than the two coexisting native species of the same functional group, Festuca arundinacea and Brachypodium phoenicoides. C. selloana reduced aboveground biomass of target native species, but not more than target native species on each other. Moreover, C. selloana did not resist interspecific competition more than target native species. Under control conditions, C. selloana did not have larger specific leaf area (SLA) and root–shoot ratio (R/S) ratio than target native species, contradicting the general statement that these traits are associated to invasiveness. F. arundinacea was the species which performed best but also the one most affected by water stress. Both C. selloana and B. phoenicoides performed in a similar way under water stress conditions. However, the alien species’ capacity to adjust to water stress, indicated by the increase in the root–shoot ratio under moderate and severe water stress, was slightly better than that of B. phoenicoides. Overall, at early recruitment stages, C. selloana is not a better competitor than the coexisting native species. However, it seems to be more resistant to water stress because as water becomes scarce C. selloana maximizes water uptake and minimizes water losses more than the native species.     

Effects of saltmarsh invasion by Spartina alterniflora on arthropod community structure and diets

Invasive plants strongly affect physical and biotic environments of native ecosystems. Insects and other arthropods as one of the major components of many ecosystems are very sensitive to subtle changes in abiotic and biotic environments. We examined the effects of exotic Spartina alterniflora invasion on community structure and diets of arthropods in a saltmarsh previously dominated by native Phragmites australis in Yangtze River estuary through net sweeping and plant harvesting methods and stable isotope analysis. Our results showed that diversity indices were not significantly different between exotic and native plant communities, but the total abundance of insects estimated through plant harvesting method was found to be lower in Spartina monoculture than that in Phragmites monoculture. Community structure of insects in Spartina monoculture was dissimilar to that in Phragmites monoculture and Phragmites–Spartina mixture. Moreover, stable carbon isotope patterns of arthropods were significantly different between Phragmites and Spartina monocultures. Although some native arthropods (perhaps generalists) shifted their diets, many native taxa did prefer Phragmites to Spartina even in Spartina monoculture. Spartina invasions resulted in reduced abundances of some arthropds, and increased dominance of others feeding preferably on Spartina. This study provides evidence that invasive plants can change the community structure and diets of native arthropods, which will eventually alter the arthropod food web, and affect the integrity and functioning of native ecosystems within a nature reserve that has been set aside for conserving the native biodiversity and maintaining the ecosystem integrity. In this sense, Spartina invasions in the Yangtze River estuary need to be managed appropriately.     

Humid tropical leguminous tree and pasture grass responsiveness to vesicular-arbuscular mycorrhizal infection

Phosphorus is the major nutrient limiting plant growth in a Costa Rican silvopastoral system located on an acid, high P-retaining, volcanic soil. We investigated plant responsiveness to vesicular-arbuscular mycorrhizal (VAM) inoculation using the leguminous tree species Erythrina berteroana Urban, and the two dominant grass species Paspalum conjugatum Berg and Homolepsis aturensis Chase of this silvopastoral system. We grew grass seedlings in the greenhouse for 15 weeks in a methyl bromide-sterilized study soil to which either mixed-species VAM inoculum (Theobroma cacao feeder roots) or autoclave-sterilized cacao roots (non-inoculated control) were added. E. berteroana was grown from both seedlings and vegetative stakes (40 cm long) for 30 and 19 weeks, respectively. Upon harvest, we measured above and below ground biomass, N and P content, rootshoot ratio, legume nodulation, and VAM infection levels. The total above-ground and root biomass of mycorrhizae-inoculated P. conjugatum seedlings were 2.5 and 2.8 times greater than those of noninoculated seedlings. In contrast, VAM-inoculated seedlings of H. aturensis produced 8.4 and 5.9 times more total above-ground and root mass than noninoculated seedlings. Mycorrhizae-inoculated E. berteroana seedlings produced 10.6 times greater shoot biomass for inoculated versus noninoculated seedlings, while E. berteroana vegetative stakes exhibited a negative growth response to VAM inoculation (an approximately 16% decrease in shoot biomass for VAM-inoculated cuttings). The difference in responsiveness between Erythrina growth forms is hypothesized to reflect the cost-benefit relationship between plant host and fungal symbiont for energy and nutrient reserves.     

Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C₃ grass, and Boutelouagracilis (HBK.) Lag., a native C₄ grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.     

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species

Summary Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.     

Evolution of an invasive phenotype: shift to belowground dominance and enhanced competitive ability in the introduced range

In response to novel selection pressures in an introduced range, non-native species may evolve more competitive phenotypes unique from those of their native range. We examined the existence of an invasive phenotype in the herbaceous perennial Artemisia vulgaris, a frequent invader of the Northeast and Mid-Atlantic US. Populations from both the native (European) and the introduced (North American) ranges were grown in intra-specific competition (same population), inter-specific competition with the native perennial herb Solidago canadensis, and alone in a common garden to quantify shifts in resource allocation and neighbor effects on performance and competitive ability. Without competition, introduced A. vulgaris populations were much shorter than native populations, but germinated earlier, produced more ramets, more belowground and total biomass, and maintained higher root-to-shoot ratios. Under inter- and intra-specific competitions, introduced A. vulgaris populations were shorter, but produced more ramets, belowground, and total biomass than native populations. S. canadensis belowground and total biomass were more highly suppressed by introduced than native A. vulgaris. Our data suggest that since the introduction to North America, A. vulgaris has evolved a more competitive invasive phenotype characterized by many short ramets with more extensive root/rhizome networks. This rapid evolutionary shift likely benefits A. vulgaris in its introduced range by allowing establishment and subsequent dominance in dense stands of existing vegetation.     

Behavioral Responses and Interactions of Estuarine Animals With an Invasive Marsh Plant: A Laboratory Analysis

Invasion by Phragmites australis into tidal marshes previously dominated by Spartina alterniflora is viewed as a serious environmental threat along the Atlantic coast of the US, but little is known about the relative habitat value of the two plants for most estuarine species. This study was designed to investigate behavioral responses, in the laboratory, of three species to the two plants. Fiddler crabs, Uca pugnax, grass shrimp, Palaemonetes pugio, and larval mummichogs (killifish), Fundulus heteroclitus were introduced into aquaria with a bare area, an area with dead Phragmites stems, and an area with dead Spartina stems. All species distributed themselves equally between the Spartina and the Phragmites. The behavior of larval mummichogs in the tanks with predators was observed. In the presence and absence of stems, they utilized the surface of the water as a refuge, as well as the stems, when present. This behavior was equally as effective as being among the stems in promoting larval survival. In microcosms with blocks of marsh with living plants, fiddler crabs and grass shrimp again did not show a preference for either species of plant, while juvenile and adult mummichogs were not consistent. Small fish chose Spartina when in the small microcosm and had no preference in the large one. Large fish chose Spartina in the small microcosm and Phragmites in the large one. Predation by adult mummichogs on grass shrimp was comparable in Spartina and Phragmites microcosms, and predation on tethered shrimp was equivalent in adjacent Spartina and Phragmites marshes in the field.     

Effects of grazing intensity on growth,reproduction, and abundance of three palatable forbs in Kansas tallgrass prairie

The effects of different intensities of cattle grazing on theaboveground growth, reproduction, and abundances of three palatable forbs werestudied in native tallgrass prairie. Populations of Asterericoides, Ruellia humilis, andAmorpha canescens were sampled at peak flowering duringthe1993–1995 growing seasons in four annually-burned sites varying incattle stocking density [ungrazed, low, moderate, high]. The threeforbs exhibited reduced shoot growth and/or reproduction under moderate toheavy grazing, and in no case did grazing increase any measure of plantperformance. Ruellia showed reduced shoot height andbiomass, percentage of stems flowering, and reproductive biomass in response tograzing. Aster showed decreases in shoot biomass andheightwith grazing. Amorpha showed no change in shoot orreproductive biomass, but a decrease in percent of flowering stems and inreproductive allocation with grazing. Patterns in the percentage of stemsgrazedindicated generally high but variable palatability among these species. Bycontrast, the three species showed inconsistent population response to grazing.Abundance (frequency) of all three species indicated no short-term changebetween years in response to grazing intensity. Responses of these speciesdiffer considerably from those of most other perennial tallgrass prairie forbsthat are unpalatable, unconsumed, and increase in performance (e.g. size,abundance) due to release from competition from the dominant grasses underungulate grazing. The results demonstrate that immediate aboveground growth andreproductive responses of established adults to grazing are not good predictorsof grazer effects on population abundances in tallgrass prairie.     

Competitive responses of the rare Viola elatior and the common Viola mirabilis

Viola mirabilis is abundant in Estonia; Viola elatior is rare. We tested whether these species differ in their competitive responses to a common grass. We used a pot experiment in which individuals of each violet species grew with 0, 2, 4, or 8 individuals of Festuca ovina in natural soil. The response patterns of shoot and root mass and leaf numbers of the two species were similar: the biomass and leaf number of both violet species decreased with increasing grass density. The decrease of root mass of the rare V. elatior was significantly greater, and the decrease of leaf number tended to be greater, than that of the common V.␣mirabilis. We conclude that the stronger competitive response of V. elatior compared to V. mirabilis might be one reason behind its lower regional and local abundance.     

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.     

两种本地植物种子萌发对飞机草的化感耐受性及其相互竞争

为探讨本地物种假地豆和白饭树对入侵植物飞机草的替代控制潜力,利用培养皿法和同质园种植实验分别研究了两个本地物种种子萌发对飞机草的化感耐受性及其与飞机草的竞争关系。结果显示:除了假地豆的萌发率在高浓度(2.5%)的飞机草叶提取液下受到显著抑制外,两个本地种的萌发在不同浓度飞机草根、茎、叶提取液下均不受抑制。飞机草与假地豆混种时,飞机草的株高、地下生物量比及根冠比显著降低,假地豆的株高无显著变化,但生物量显著增加;飞机草的竞争参数相对产量(RY)显著小于1,竞争攻击力系数显著小于零,表明其竞争力弱于假地豆。飞机草与白饭树混种时,飞机草的根冠比也显著降低,但株高和生物量均显著增加,而白饭树的株高和生物量却显著降低;飞机草的相对产量(RY)显著大于1,竞争攻击力系数显著大于零,表明其竞争力强于白饭树。结合以上结果,本地植物假地豆可以一定程度上竞争抑制飞机草的生长,具有替代控制飞机草的潜力,而白饭树可以在清除飞机草后的入侵地辅助植被修复。实验结果为飞机草替代控制的目标物种的筛选提供了科学依据,对被飞机草入侵生境生物多样性的恢复和重建具有一定指导意义。