California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Arthropod diversity of exotic vs. native Robinia species in northern Arizona

1 Arthropods were collected on native locust, Robinia neomexicana A. Gray, and exotic Robinia pseudoacacia L. in northern Arizona over a 2‐year period to determine the number of arthropod species and number of individuals present. 2 More arthropod species were found on the native (251) than on the exotic Robinia (174). 3 Greater species diversity was likewise found on the native than the exotic. The five most numerous insects collected each year accounted for 81% to 91% of the total number collected on the exotic and native Robinia in 1997 and 1998. Only 12 species occurred on both the native and exotic Robinia in both years. 4 These findings are discussed in the context of using exotic trees in plantations and ecological theory regarding rates of arthropod species accumulation on exotic hosts.     

The response of native species to removal of invasive exotic grasses in a seasonally dry Hawaiian woodland

Abstract. Non-native perennial grasses form 30% of the live understory biomass in seasonally dry, submontane forests in Hawaii Volcanoes National Park, yet their effects on native species are unknown. We removed these grasses from plots of 20 m × 20 m in 1991 and maintained removal and control areas over the next three years. Two fast growing shrub species, Dodonaea viscosa and Osteomeles anthylidifolia, increased in size significantly more in removal areas than in controls. Individuals of the most abundant shrub species, Styphelia tameiameia showed no net growth response to grass removal. They did, however, change their architecture: many branches along the mid and upper sections of the main trunk died and a proliferation of new leaves and shoots occurred in the lower 40 cm of trunk. Basal diameter increase was very small in Metrosideros polymorpha, the dominant tree species in these sites. All species except Styphelia had significantly increased leaf tissue nitrogen in removal plots by 18 months after removal when compared to shrubs in control areas suggesting that removal plot shrubs had greater access to soil nitrogen. Available soil-N pools, which were generally higher in the removal plots, support this interpretation. Light levels near the soil surface were also higher where grasses were removed than where they were present which may have contributed to increased shrub growth. By contrast, soil moisture was consistently lower where grasses were removed than where they were still present. Shrub tissue carbon isotope values were consistent with the interpretation that shrubs in removal plots had less rather than more water available to them. Hence, the increased growth observed in removal plot shrubs could not be due to release from moisture competition. Lastly, our results showed that seedlings of all woody species except Metrosideros were significantly more abundant in removal plots at both one and three years after removal and initially high sapling mortality was balanced by high recruitment into the sapling class. We believe that over time this will result in increased densities of native shrubs if grasses are kept out. With the presence of grasses, shrub growth in these woodlands is reduced and biomass is shifting towards grasses.     

Coexistence and interference between a native perennial grass and non-native annual grasses in California

Little is known about the potential for coexistence between native and non-native plants after large-scale biological invasions. Using the example of native perennial bunchgrasses and non-native annual grasses in California grasslands, we sought to determine the effects of interference from non-native grasses on the different life stages of the native perennial bunchgrass Nassella pulchra. Further, we asked whether N. pulchra interferes with non-native annual grasses, and whether competition for water is an important component of these interspecific interactions in this water-limited system. In a series of field and greenhouse experiments employing neighbor removals and additions of water, we found that seedling recruitment of N. pulchra was strongly seed-limited. In both field and greenhouse, natural recruitment of N. pulchra seedlings from grassland soil was extremely low. In field plots where we added seeds, addition of water to field plots increased density of N. pulchra seedlings by 88% and increased total aboveground N. pulchra seedling biomass by almost 90%, suggesting that water was the primary limiting resource. In the greenhouse, simulated drought early in the growing season had a greater negative effect on the biomass of annual seedlings than on the seedlings of N. pulchra. In the field, presence of annuals reduced growth and seed production of all sizes of N. pulchra, and these effects did not decrease as N. pulchra individuals increased in size. These negative effects appeared to be due to competition for water, because N. pulchra plants showed less negative pre-dawn leaf water potentials when annual neighbors were removed. Also, simply adding water caused the same increases in aboveground biomass and seed production of N. pulchra plants as removing all annual neighbors. We found no evidence that established N. pulchra plants were able to suppress non-native annual grasses. Removing large N. pulchra individuals did not affect peak biomass per unit area of annuals. We conclude that effects of interference from non native annuals are important through all life stages of the native perennial N. pulchra. Our results suggest that persistence of native bunchgrasses may be enhanced by greater mortality of annual than perennial seedlings during drought, and possibly by reduced competition for water in wet years because of increased resource availability. Received: 12 November 1998 / Accepted: 4 August 1999     

本地种与外来种水生植物在不同基质营养下的生长比较

近年来,生物入侵由于其对生态系统,环境,经济等多方面造成的后果日益严重,逐渐成为目前全球普遍关注的生态问题。对于植物而言,外来入侵种不仅会大大降低本地物种的存活率,而且对本地物种的分布也会造成很大程度的负面影响。不仅如此,外来入侵种还会对本地水生或陆生生态系统造成十分严重的影响和破坏。随着国内外专家学者对外来种研究的深入,许多有关外来种入侵机制的假说和理论不断被提出。其中以资源假说(Resource Hypothesis,Davis 等 2000)研究的最为广泛和深入。在资源假说中,Davis等提出:伴随着一定数量繁殖体,外来物种的入侵会随着资源(例如光,营养和水份)的增加而加剧。为了验证(1)外来种和本地种的表现是否存在差异,如果存在,这种是否也会随着资源的增加而增强?(2)外来种与本地种的差异是否对我们预测和防治入侵有帮助?本文在此基础上,采用4种水生外来种,水鳖科(Hydrocharitaceae)两种,小二仙草科(Haloragidaceae)狐尾藻属(Myriophyllum)两种,与本地同科或同属且形态相似的3种水生植物为实验材料。分别配制两种不同营养级别的基质环境,比较了在高、低营养基质环境中外来种和本地种与生长相关的参数指标。结果显示:在高营养基质和低营养基质环境中,本地种较外来种更大地积累其生物量并且拥有更高的相对生长速率(relative growth rate)。结果还显示:在侧枝的生物量分配方面,外来种较本地种显著增高。由于断枝是水生植物无性繁殖的重要手段之一,并且断枝由侧枝或主茎受到生物或非生物的扰动产生。我们使用侧枝生物量的分配这一指标来间接反映水生植物的无性繁殖能力。结果表明,本地种较外来种能较好适应当地不同的基质营养环境;但是在无性繁殖体的传播方面,外来种较本地种占有一定的优势。因此,我们尝试提出:对外来水生植物的防治应继续以管理本地水生植物的栖息地和保护本地水生植物多样性为主,并且应加大对外来水生植物无性繁殖体特别是断枝传播的管理和控制力度。     

Competition for nitrogen between Australian native grasses and the introduced weed Nassella trichotoma

BACKGROUND: and Aims Nassella trichotoma is an unpalatable perennial grass weed that invades disturbed native grasslands in temperate regions of south-eastern Australia. This experiment investigated whether elevated N levels, often associated with disturbance, increases the competitiveness of N. trichotoma relative to C3 and C4 native Australian grasses. METHODS: A pot experiment investigated competitive interactions between four native grasses, two C3 species (Microlaena stipoides and Austrodanthonia racemosa) and two C4 species (Themeda australis and Bothriochloa macra), and N. trichotoma at three different N levels (equivalent to 0, 60 and 120 kg ha-1) and three competing densities (zero, one and eight neighbouring plants), using an additive design. KEY RESULTS: All native grasses were competitive with N. trichotoma at low N levels, but only M. stipoides was competitive at high N. High densities of native grasses (8:1) had a major competitive effect on N. trichotoma at all N levels. The competitive ranking of native grasses, across all N levels, on N. trichotoma was: M. stipoides>A. racemosa>B. macra>T. australis. The C3 species were generally more competitive than the C4 species and C4 grasses were not inherently more productive at low N levels, in contrast to the results of other studies. CONCLUSION: To resist invasion from N. trichotoma, these native grasses need to be maintained at a high density and/or biomass. The results do not support the theory that species such as N. trichotoma, with high tissues density, are always less competitive than those of low tissue density; in this case competitiveness depended on N levels. The ability of N. trichotoma to accumulate biomass at a higher rate than these native grasses, helps to explain why it is a major weed in disturbed Australian native grasslands.     

火与外来植物相互关系的研究进展

入侵种对本地的生态系统和生物多样性均有不良的影响,严重的会造成物种的灭绝和生态系统的崩溃,这已在全球范围引起广泛关注.在植物外来种与火生态因子的作用研究中发现,火与外来种的关系随物种生物学特性、火作用的时间、频度、强度不同而不同,火有时会有效地抑制外来种的生长和入侵,有时会促进一些外来种的生长和入侵.反之,一些外来种会对火的产生起到积极的作用,一些外来种又会抑制火的发生.火作为控制入侵种的一种方法,经科学地运用,可对某些入侵种起到有效的控制作用.     

Species diversity in native fish community in Japan: comparison between non-invaded and invaded ponds by exotic fish

The relationship between invasions by two exotic fishes (Micropterus salmoides and Lepomis macrochirus) and species diversity in native fish communities was studied in 14 Japanese farm ponds. We found that mean number of species in native fish communities was three times higher in the ponds without the exotic fish than in the ponds with them. Further, negative relationships were observed between abundance of the two exotic fish and the total abundance of native fish communities. Our results suggest that invasions by the two exotic fish caused serious depletion of native fish communities, although another process can also be considered , that is, that ponds with poor native fish communities were prone to colonization by these exotic fish.     

Effect of nitrogen, water and neighbor density on the growth of Hesperis matronalis and two native perennials

Disturbance is a well-recognized catalyst of exotic species invasion, depriving or releasing resources into communities and favoring the spread of some invaders. Hesperis matronalis (dame’s rocket) is widespread in North America and has potential to become a major problem in natural communities due to anthropogenic influences. We used a target-neighbor design in a greenhouse to grow H. matronalis and native seedlings (Campanula rotundifolia and Muhlenbergia montana) at various levels of neighbor density, nitrogen and water. H. matronalis clearly reduced C. rotundifolia and M. montana aboveground growth and maintained its competitive advantage across all treatments. We expected H. matronalis to maximize its growth and have the greatest negative effect on native species under high resource conditions because ruderal species are able to take advantage of excess resources. H. matronalis demonstrated a clearly negative effect on native species, but a particular ability to outcompete native species at high resource levels depended on resource and native species identity. We also expected that the native species would not exploit excess resources as well as H. matronalis, instead growing better under ambient or low resource conditions because of local adaptation. Rather, M. montana benefited from high water inputs, inconsistent with plant strategies characteristic of stress tolerators. Information on the effects of H. matronalis on native plants in a controlled setting may aid land managers to understand its potential effects in natural communities.     

The alien flora of Europe: a taxonomic and biogeographic review

Abstract. A geographic and taxonomic overview of the non-indigenous plant species of Europe, based on the ‘Flora Europaea’ is given. The flora of Europe includes 1568 species which have either expanded their ranges within Europe under human influence (naturalized European species) or are of non-European origin (exotic species). The latter group consists of 580 species (37%) which form a diverse group in terms of their taxonomic composition and geographic origin. The exotics are represented by 113 families, of which the Compositae, Rosaceae and Gramineae are most important. The ratio of species to families is low. Most exotic species in Europe originate from the Americas and Asia. Countries of southern Europe have a higher relative number of exotics in their non-native flora than northern ones. The species-range size distribution differs between naturalized European and exotic species; the latter are on average more widespread than the naturalized.     

Composition of soil seed banks in southern California coastal sage scrub and adjacent exotic grassland

Soil seed banks are important to many plant communities and are recognized as an important component of management plans. Understanding seed bank composition and density is especially important when communities have been invaded by exotic species and must be managed to promote desirable species. We examined germinable soil seed banks in southern California coastal sage scrub (CSS) that is heavily invaded by exotic grasses and in adjacent exotic grassland. Soils from both communities had similar seed banks, dominated by high densities of exotic grass and forb species. Up to 4,000 exotic grass seeds and at least 400 exotic forb seeds/m² were found in most soils, regardless of aboveground vegetation type. Native forbs averaged 400 seeds/m² in grass-dominated areas and about 800 in shrub-dominated soils. Shrub seed density was <1 and <10 seeds/m² in grass- and shrub-dominated areas, respectively, indicating that the shrub seed bank is not persistent compared to annuals. We also compared pre- and post-burn soil seed banks from one location that burned in October 2003. Late-season burning in both grass- and CSS-dominated areas disproportionately reduced exotic grass seed densities relative to native seed densities. The similarity of the seed banks in adjacent grass and shrub communities suggests that without intervention, areas currently dominated by CSS may become more similar to grass-dominated areas in terms of aboveground vegetation. In such areas, the first growing season following a wildfire is a window of opportunity for increasing native diversity at a time when density of exotic grass seeds is low. At time of research, Robert D. Cox was graduate student.     

Differential responses to nitrogen fertilization in native shrubs and exotic annuals common to mediterranean coastal sage scrub of California

This study examined the growth responses of exotic annuals and native shrubs to elevated N levels to test the hypothesis that increased N availability favors nitrophilous annuals over the slower-growing shrubs. The vegetation structure of the coastal sage scrub ecosystems in southern California is shifting from shrubland to annual grasslands. Over the last 30 years large tracts of wildlands, particularly those adjacent to urban centers, have lost significant native shrub cover, which has been replaced by exotic annuals native to the Mediterranean Basin. During this same time, air pollution has led to increased terrestrial eutrophication by atmospheric deposition. Changes in vegetation are often the result of changes in resource availability. The results of our experiments showed the three native shrubs tested to be more nitrophilous than the three annuals tested, which contrasts with most models of perennial species' adaptation to stressful environments. Under greenhouse conditions the annual grasses exhibited yield depression at the highest N treatments of 80 g g^–1 in soil. The three shrub species evaluated continued to increase shoot biomass at 80 g g^–1 N in soil. The grasses also exhibited increased tissue N concentrations with increased soil N in contrast with the shrubs where there was little difference in tissue N concentrations with increasing availability. Although the differential yield responses to elevated N do not explain the success of the annual vegetation in replacing shrubs, the inability of the shrubs to regulate growth under elevated N levels may explain the poor survival of mature individuals.     

Effects of elevated CO2 on competition between native and invasive grasses

Oecologia - Elevated atmospheric CO2 concentration increases the performance of invasive plants relative to natives when grown in monoculture, but it is unclear how that will affect the relative...     

Capture and allocation of nitrogen byQuercus douglasii seedlings in competition with annual and perennial grasses

Summary The spatial overlap of woody plant root systems and that of annual or perennial grasses promotes competition for soil-derived resources. In this study we examined competition for soil nitrogen between blue oak seedlings and either the annual grassBromus mollis or the perennial grassStipa pulchra under controlled outdoor conditions. Short-term nitrogen competition was quantified by injecting¹⁵N at 30 cm depth in a plane horizontal to oak seedling roots and that of their neighbors, and calculating¹⁵N uptake rates, pool sizes and¹⁵N allocation patterns 24 h after labelling. Simultaneously, integrative nitrogen competition was quantified by examining total nitrogen capture, total nitrogen pools and total nitrogen allocation.Stipa neighbors reduced inorganic soil nitrogen content to a greater extent than didBromus plants. Blue oak seedlings responded to lower soil nitrogen content by allocating lower amounts of nitrogen per unit of biomass producing higher root length densities and reducing the nitrogen content of root tissue. In addition, blue oak seedlings growing with the perennial grass exhibited greater rates of¹⁵N uptake, on a root mass basis, compensating for higher soil nitrogen competition inStipa neighborhoods. Our findings suggest that while oak seedlings have lower rates of nitrogen capture than herbaceous neighbors, oak seedlings exhibit significant changes in nitrogen allocation and nitrogen uptake rates which may offset the competitive effect annual or perennial grasses have on soil nitrogen content.     

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments. Laurel Pfeifer-Meister and Esther M. Cole contributed equally to this work.     

The roles of exotic grasses and forbs when restoring native species to highly invaded southern California annual grassland

Many semi-arid shrublands in the western US have experienced invasion by a suite of exotic grasses and forbs that have altered community structure and function. The effect of the exotic grasses in this area has been studied, but little is known about how exotic forbs influence the plant community. A 3-year experiment in southern California coastal sage scrub (CSS) now dominated by exotic grasses was done to investigate the influence of both exotic grasses (mainly Bromus spp.) and exotic forbs (mainly Erodium spp.) on a restoration seeding (9 species, including grasses, forbs, and shrubs). Experimental plots were weeded to remove one, both, or neither group of exotic species and seeded at a high rate with a mix of native species. Abundance of all species varied with precipitation levels, but seeded species established best when both groups of exotic species were removed. The removal of exotic grasses resulted in an increase in exotic and native forb cover, while removal of exotic forbs led to an increase in exotic grass cover and, at least in one year, a decrease in native forb cover. In former CSS now converted to exotic annual grassland, a competitive hierarchy between exotic grasses and forbs may prevent native forbs from more fully occupying the habitat when either group of exotics is removed. This apparent competitive hierarchy may interact with yearly variation in precipitation levels to limit restoration seedings of CSS/exotic grassland communities. Therefore, management of CSS and exotic grassland in southern California and similar areas must consider control of both exotic grasses and forbs when restoration is attempted.     

Demographic changes and the development of competition in a native Australian eucalypt forest invaded by exotic pines

Data from the second half of a 14-year study of a eucalypt dry sclerophyll forest invaded by Pinus radiata D. Don shows further progress in the development of a mixed eucalypt-pine stand. Earlier (Chilvers and Burdon 1983) it was clear that pines invading from an adjacent commercial plantation were starting to contribute their own progeny to the mixed stand. Initially the recruitment of established pines (> 1 m high) from these seedlings was very slow. However, since 1984, increasing numbers of seedlings have survived beyond their first year, suggesting the possibility of another wave of successful establishment in the near future. The established fraction of the pine population continued to grow rapidly and incurred no mortality. Many of these plants have now emerged through the top of the eucalypt canopy. In contrast, both juvenile and adult eucalypt numbers continued to decline. Despite these demographic and visual changes, comparative analysis of the growth of established pines over two consecutive 7-year periods (1974–1981; 1981–1988) showed that competition between neighbouring plants, especially pines, is beginning to affect individual growth rates. Growth rates of pines having other pines as nearest neighbours declined 40% between 1974–1981 and 1981–1988, while that of pines with eucalypts as nearest neighbours fell by 28%. In pine-pine nearest neighbour (NN) pairs, growth rates were significantly affected by the size (diameter) of the nearest neighbour. In pine-eucalypt NN pairs growth rates of the pines were not affected by the size of the eucalypt, but these were significantly negatively correlated with the inverse of the distance to the eucalypt. More broad-based assessment of 43 pines showed a significant relationship between their growth rates and the summed competitive effect of other pines within a 5-m radius. No such relationship was found between growth rate and the summed competitive effect of eucalypts in the 5-m zone.