Management regimes for a plant invader differentially impact resident communities

Management of plant invaders causes disturbance and the compounding effects of both invasion and management on biodiversity merit rigorous assessment. We explored the impacts of both plant invasion and management disturbance in coastal dune communities of Australia invaded by the introduced shrub bitou bush (Chrysanthemoides monilifera ssp. rotundata). We compared the effects of intensive, manually based invader control with those of extensive control (mainly herbicide spraying from aircraft) across fore dune communities. We examined one management regime, which was typically intensive, in hind dune communities. Our main aim was to test the prediction that intensive methods would achieve better biodiversity outcomes than extensive approaches in fore dune communities. We found that extensive management created native species complements that diverged from non-invaded sites while intensive management approached non-invaded site conditions. However, intensively managed sites were also richer in other exotic species than extensively managed sites. Thus, while biodiversity outcomes were better under intensive management, the risk profile of such sites was increased by the greater array of exotics that could potentially replace the original invader. The positive effects of management evident on some fore dunes were not found in hind dunes, where managed sites had the highest exotic species richness and similar bitou bush seedling abundance as both non-invaded and bitou-invaded sites. Hind dune managed sites were also compositionally distinct from non-invaded sites. Thus fore and hind dune communities exhibited different responses to bitou management regimes.     

Soil seed bank compositional change constrains biodiversity in an invaded species-rich woodland

Relationships between plant invasion and the soil seed bank in highly diverse fire adapted mediterranean woodlands are poorly understood, yet critical for that ecosystem’s conservation. Within the biodiversity hotspot of southwest Australia we investigated the composition and diversity of the Banksia woodland soil seed bank in good condition (GC), medium condition (MC) and poor condition invaded by the South African perennial species Ehrharta calycina (PCe) and Pelargonium capitatum (PCp). The investigation assessed three questions: (1) Do soil seed banks of invaded sites have fewer germinants of native species and more germinants of introduced (non-native) species than sites with minimal invasion? (2) Do soil seed banks show shifts in ecological functional types with invasion? (3) Is the soil seed bank of introduced species persistent? Native species germinants, mainly shrubs and perennial herbs, were highest in GC sites and least in poorer condition sites suggesting a reduction in their numbers had occurred over time. Introduced germinants were dominated by perennial and annual grasses, and annual herbs. E. calycina had the greatest seed density (8328 germinants m⁻²). More introduced than native germinants occurred in the litter. Rapid germination of introduced species (30% in week 1) compared to native species (4% in week 1) provides the capacity for their early dominance. A limited native soil seed bank and dominant persistent introduced soil seed bank represent great challenges for the structural and functional conservation and restoration of woodland ecosystems. This study provides key new knowledge, applicable to a wide range of ecosystems, to help formulate conservation protocols to control dominant introduced species and conserve and restore biodiverse-rich woodlands.     

Exotic woody invader limits the recruitment of three indigenous plant species

The demographic response of indigenous plants to the invasion of exotic woody plants has rarely been quantified, however, could be beneficial to restoration efforts. We determined which life history stages of three indigenous plants: Correa alba var. alba (Rutaceae), Monotoca elliptica (Epacridaceae) and Lomandra longifolia (Lomandraceae), were most affected by the invasion of Chrysanthemoides monilifera spp. rotundata (bitou bush) on the eastern Australian coast. We also assessed whether various morphological and physiological parameters of the mature stage of these species were affected by the presence of bitou bush. Populations of all three indigenous species in bitou bush invaded habitats had significantly fewer small individuals and lower population density than those in non-invaded habitats. The mean flower production, number of vegetative buds, and physiological stress of the mature stage of each of these species in bitou bush invaded habitat did not differ from those present in the non-invaded habitat. We therefore propose that bitou bush affects indigenous plant populations primarily by preventing recruitment through the germination or seedling growth stages. The reduction in indigenous plant recruitment creates spaces that are likely to facilitate bitou bush monoculture formation in the new host environment. Planting of established juvenile plants is suggested to assist in the restoration of bitou bush invaded areas.     

Herbicides, weeds and endangered species: management of bitou bush (Chrysanthemoides monilifera ssp. rotundata) with glyphosate and impacts on the endangered shrub, Pimelea spicata

Environmental weed invasion threatens the biodiversity of native species. Unfortunately, managing these weeds may also affect biodiversity adversely. A recent example occurred when glyphosate, a herbicide used to control the highly invasive weed, bitou bush (Chrysanthemoides monilifera ssp. rotundata), accidentally drifted over a small population of an endangered shrub, Pimelea spicata. Following concerns that the affected population would not recover and, thereby, cause the local extinction of P. spicata, we conducted a series of glasshouse and field experiments to explore the impacts of glyphosate on this endangered species. Seedlings and young plants of P. spicata, in which the tap root was undeveloped, were killed by a single application of glyphosate. Older plants with a well developed tap root also died back initially, but about 50% of individuals re-sprouted. This re-growth was associated with a significant decrease in tap root diameter, implying that further disturbance, including repeated treatment with glyphosate, would kill plants by impairing their potential for recovery. Unlike some sclerophyllous native shrubs, the tolerance of P. spicata to glyphosate was limited, even when its growth was slowed artificially by limiting water availability. Winter applications of glyphosate to manage infestations of bitou bush will impact adversely on populations of P. spicata and may also affect the other rare and endangered species whose survival is threatened by this species, even though some natives are unaffected by the herbicide. Protecting native biodiversity from bitou bush will involve sustainable weed management that minimises impacts on non-target native species.     

The effects of invasive North American beavers on riparian plant communities in Cape Horn, Chile: Do exotic beavers engineer differently in sub-Antarctic ecosystems?

North American beavers (Castor canadensis) were introduced into southern South America in 1946. Since that time, their populations have greatly expanded. In their native range, beavers shape riparian ecosystems by selectively feeding on particular plant species, increasing herbaceous richness and creating a distinct plant community. To test their effects as exotic engineers on sub-Antarctic vegetation, we quantified beaver impacts on tree canopy cover and seedling abundance and composition, as well as their impacts on herbaceous species richness, abundance and composition on Navarino Island, Cape Horn County, Chile (55°S). Beavers significantly reduced forest canopy up to 30 m away from streams, essentially eliminating riparian forests. The tree seedling bank was greatly reduced and seedling species composition was changed by suppressing Nothofagus betuloides and Nothofagus pumilio, but allowing Nothofagus antarctica. Herbaceous richness and abundance almost doubled in meadows. However, unlike beaver effects on North American herbaceous plant communities, much of this richness was due to invasion by exotic plants, and beaver modifications of the meadow vegetation assemblage did not result in a significantly different community, compared to forests. Overall, 42% of plant species were shared between both habitat types. Our results indicate that, as predicted from North American studies, beaver-engineering increased local herbaceous richness. Unlike in their native range, though, they did not create a unique plant community in sub-Antarctic landscapes. Plus, the elimination of Nothofagus forests and their seedling bank and the creation of invasion pathways for exotic plants together threaten one of the world’s most pristine temperate forest ecosystems.     

Distribution, species composition and management implications of seed banks in southern New England coastal plain ponds

Buried seeds that germinate during periods of low water or water level drawdown can play important roles in shaping plant community composition, community dynamics and species richness in ecosystems with fluctuating water levels. Northeastern US coastal plain ponds have fluctuating water levels and contain a characteristic shoreline flora that contains many rare plants. The objectives of this study were to: (1) test whether geographically distant ponds in Cape Cod and Martha’s Vineyard had distinct seed banks, (2) determine if hydrologic status as permanent and ephemeral ponds led to differences in seed banks, and (3) examine seed diversity and seed abundance across gradients of shoreline elevations and sediment characteristics. Viable seeds of 45 plant species were identified from nine ponds. Native species dominated pond-shore seed banks and made up 89-100% of all species. There was high overlap in seed bank composition across hydrological classes and geographic regions. One hydrological class captured 73-76% of total species and one geographical region captured 69-78% of the total species recovered from the entire suite of seed bank samples. Seeds were relatively evenly distributed along the shorelines of ephemeral ponds but seed diversity and abundance were lower at low elevations in permanent ponds. Results suggest that strategies to protect pond shorelines to capture maximum diversity of coastal plain pond plants contained in pond sediment seed banks should be implemented across pond hydrologic classes and across a wide geographic area. Shoreline seed distributions indicate that ground-water withdrawals or climate changes that lower pond water levels in permanent ponds will reduce the diversity and abundance of plants recovered from seed banks by shifting water levels to a shoreline zone of high sediment organic matter where seed densities are lower. This effect will be much less in ephemeral ponds where seed diversity and abundance on pond bottoms was high.     

黄土丘陵沟壑区红砂灌丛土壤种子库及其自然更新潜力评估

[目的]对黄土丘陵沟壑区红砂灌丛植被土壤种子库的特征及红砂灌丛植被自然更新潜力进行分析和评估,以说明红砂灌丛土壤种子库在植被恢复与重建中的重要作用。[方法]采用土壤种子库"萌发法"(每个地段10个2m×2m样方内分表土层0—2cm和2—5cm土层采集土样)及野外植被调查方法,对黄土丘陵沟壑不同坡位红砂灌丛土壤种子库进行研究。[结果]黄土丘陵沟壑区红砂灌丛植被土壤种子库发芽试验共观察到的4 251株幼苗,分属于9个物种。土壤种子库密度在100~1 000粒/m2,物种数在0.8~1.2种/0.01m2,下坡段的土壤种子库平均种子密度和平均物种数均比上坡段高。3个坡段的2个层次的平均种子密度和平均物种数均随土层加深而减小。3个坡段土壤种子库和地上植被的组成物种多为草本植物和红砂灌丛,地上植被与其土壤种子库的密度及物种数均呈不显著相关;物种组成的Sorensen相似性指数较高,均达到0.60以上,且土壤种子库比地上植被具有更高的物种丰富度。[结论]黄土丘陵沟壑区红砂灌丛具有依靠土壤种子库实现自然更新的潜力,但由于物种组成种类较少,现存灌丛植被一旦遭到破坏,仅靠土壤种子库恢复现存植被是困难的,需要在对自然恢复潜力评估的基础上,积极采取人工保护促进自然恢复的策略。     

Evaluating barriers to native seedling establishment in an invaded Hawaiian lowland wet forest

Many tropical island forest ecosystems are dominated by non-native plant species and lack native species regeneration in the understorey. Comparison of replicated control and removal plots offers an opportunity to examine not only invasive species impacts but also the restoration potential of native species. In lowland Hawaiian wet forests little is known about native species seed dynamics, recruitment requirements, or the effects of management. In a heavily invaded lowland wet forest, we examined the relationship between seed presence and seedling establishment in control and removal plots. Non-native species were competitively superior because they had higher germination percentages and dominated the seed bank; only seven out of 33,375 seedlings were native. In contrast, the seed rain contained native seed, but native seedling recruitment was almost exclusively limited to removal plots, suggesting that optimum establishment conditions are not met in the presence of a dense mid-storey of non-native species. Non-native species dominance was altered and biomass significantly decreased over time resulting in a reduced weeding effort (12.38-0.77 g day⁻¹). We suggest that with opening of the canopy through non-native species removal and subsequent weeding, it may be possible to reduce the seed bank enough to skew the regeneration potential towards native species. Our results suggest that germination success and lack of a seed bank are the main bottlenecks for native species. We conclude that without invasive species control, future regeneration of Hawaiian lowland wet forests is likely to be almost entirely non-native.     

The Vertical Distribution of Soil Seed Bank and Its Restoration Implication in an Active Sand Dune of Northeastern Inner Mongolia,China

Although the functions and characteristics of soil seed banks in topsoil layers have been described for various ecosystems, the spatiotemporal pattern of the seed bank in deep soil and its ecological implications for vegetation restoration of active sand dune have not been fully explored. In 2007 and 2008, seed densities with regard to dune position, soil depth and season were investigated on an active and a stabilized sand dune of northeastern Inner Mongolia, China. Seeds in the 0–10 cm topsoil layer accounted for 60% of total soil seed bank on the stabilized sand dune, while more than 40% of seeds were stored in the 50–100 cm layers on the active sand dune. Seed density declined significantly with soil depth on the stabilized sand dune, but it was relatively constant across the 0–100 cm soil profile on the active sand dune. Seed density fluctuated with soil depth on the active sand dune suggesting that seeds were either relocated upward or downward over time. Seeds of annual non‐psammophytic species accounted for the majority of soil seed bank on the stabilized sand dune, while pioneer psammophytes contributed more to the soil seed bank of the active sand dune. Our data suggest that seeds in the deep soil layers of active sand dunes account for a large proportion of the whole soil seed bank. Because of the effect of wind erosion, seeds in deep soil could be gradually exposed to shallow soil layers and potentially contribute to population recruitment and vegetation restoration on active sand dunes. Copyright © 2015 John Wiley & Sons, Ltd.     

子午岭天然柴松林土壤种子库

采取子午岭天然柴松林地3个坡位与1个对照区共4个大样地54个小样方的土壤样品,通过室内种子萌发和地上植被调查相结合的方法对其种子库进行研究,以揭示其土壤种子库特征及与地上植被的关系。结果显示：1）土壤种子密度平均为1 496.7粒/m^2,最小为1 240.0粒/m^2,最大为1 785.0粒/m^2,荒山对照区为510.0粒/m^2;2）种子库的垂直分布格局显示,近90%的种子储存于枯枝落叶层和0-7.5 cm土层中;3）研究中共统计到植物种32种,分属于20科,其中多年生草本和常绿灌木种子比例最高,分别占种子库种子总数的48.3%和41.9%;4）不同坡位的物种丰富度指数介于2.523-3.082之间,而物种多样性指数介于1.458-1.781之间,其中中坡物种丰富度指数最大而多样性指数最小,上坡多样性指数和均匀度指数都最大;5）各坡位地上植被主要建群植物种中,除白颖苔草、大披针苔草、胡枝子、唐松草在对应坡位的土壤种子库中出现且分别占该坡位种子库的19.7%、2.1%和0.8%、1.7%和3.1%0、.4%外,其他建群种如榛子、子、地榆等在种子库中均未出现,种子库不能很好地反映地上植被组成,部分未出现物种与其自身萌发性质有关;6）土壤种子库中柴松种子所占比例约为0.6%（地上植被调查发现）,柴松林下有更新幼苗,约5-7粒/m^2,说明种子库具有潜在萌发能力,森林具备一定的自我更新能力。     

The feasibility of using soil seed bank for natural regeneration of degraded sandy grasslands

Desertification in degraded grasslands is manifested through the development of bare sandy patches, which eventually lead to habitat fragmentation. The ability of these bare sandy patches to regenerate naturally through in-situ soil seed banks is not well understood. To fill this knowledge gap, we randomly selected 24 bare sandy patches with areas ranging from 19 to 898 m² in a desertified grassland of the Horqin sandy land, Northern China to determine whether soil seed bank can be used for natural regeneration of bare sandy patches. Species composition and density of soil seed bank as well as aboveground vegetation composition, abundance and coverage were investigated. We then determined their relationships with in-situ habitat characteristics. Our observations showed that the studied area had low soil seed bank density and species richness, as well as depauperate soil seed bank communities. Consequently, local soil seed bank was not able to provide sufficient seed source for natural regeneration. This was indicated by the relationships between aboveground vegetation, soil seed bank and the in-situ habitat characteristics. For bare patches with an area between 300 m² and 900 m², increase the soil seed bank density and species richness should be the main restoration measures. For bare patches with a small area of less than 50 m², restoration of vegetation density should be the main measure. Our data highlighted that different extents of desertification, indicated by different bare patches, are requiring distinct restoration measures.     

Impact threshold for an alien plant invader, Lantana camara L., on native plant communities

Alien plant invaders significantly threaten native community diversity, although it is poorly understood whether invasion initiates a linear or non-linear loss of resident species. Where low abundances of an invader have little impact on native species diversity, then a threshold level may exist, above which native communities rapidly decline. Our aim was to assess the broadscale effects of an alien thicket-forming shrub, lantana (Lantana camara L.), on wet sclerophyll forest in southeastern Australia. Vascular plant species richness, abundance and composition were measured and compared along a gradient of lantana invasion. There was a strong negative non-linear relationship between native species richness and lantana cover, indicative of an impact threshold. Native species richness remained stable below 75% lantana cover, but declined rapidly above this threshold level, leading to compositional change. Thus, sparse lantana infestations had evidently little effect on the resident community, with impacts elicited at an advanced stage of invasion. The impact of lantana was pervasive, with all major structural groups (i.e. ferns, herbs, shrubs, trees and vines) exhibiting significant species losses; however, the rate of species loss was relatively greater for tree and shrub species, signalling a shift in vegetation structure from tall open forest to low, dense lantana-dominated shrubland. Potentially, broadscale conservation of species diversity could be achieved by maintaining lantana infestations below the 75% cover impact threshold at sites containing regionally common species that are also widely represented in non-invaded vegetation. This would enable targeted invader eradication at sites of high conservation value (i.e. those containing regionally rare species or endangered ecological communities).     

Diversity declines in Microstegium vimineum (Japanese stiltgrass) patches

The spread of invasive plant species and their impacts on plant communities have received international attention as global trade and global environmental change enhance the colonization and establishment of introduced species and threaten the integrity of native ecosystems. Because introduced species vary in their impact, studying the relationship between invasion and native communities is necessary to guide allocation of finite management resources. By studying adjacent pairs of invaded and uninvaded plots across an eastern United States forested landscape, we demonstrate Microstegium vimineum was associated with local declines in species richness and cover of native species. Negative impacts of M. vimineum on species richness did not emerge until August when M. vimineum cover and height were greatest, highlighting the value of following study subjects through the growing season. In contrast, native species cover was already lower in invaded plots early in the growing season. M. vimineum invasion was not the only important driver of species richness and community composition within the study region; abiotic environmental gradients, such as soil nitrate concentration and pH, across the six study sites were also important in affecting species richness and cover, but lessened in explanatory power through the growing season. We conclude that M. vimineum has effects on community structure that may have long-term consequences for biodiversity. Studies which track sites through time and consider multiple scales are required as invaders impact multiple biotic and abiotic factors operating at different spatial and temporal scales.     

SOIL SEED BANK CHARACTERISTICS BENEATH AN AGE SEQUENCE OF CARAGANA MICROPHYLLA SHRUBS IN THE HORQIN SANDY LAND REGION OF NORTHEASTERN CHINA

To understand how shrub cover affects the spatiotemporal patterns of the soil seed bank and to assess the role of vegetation in the restoration of desertified land in a semi‐arid region of China, we investigated the species composition and seed density of the soil seed bank under and outside the canopies over two seasons for an age sequence of Caragana microphylla shrubs in the Horqin Sandy Land region, Inner Mongolia. The results showed that a total of 24 plant species seeds were present in the soil seed bank, of which 20 were annuals or biennials. The seed densities in the soil seed bank were in the range of 830 – 13882·5 seeds m⁻² at 10 cm depth. Species richness in the soil seed bank did not increase as the shrubs aged, whereas the seed densities increased significantly. Five annual species: Setaria viridis, Eragrostis pilosa, Chenopodium acuminatum, Chenopodium glaucum and Corispermum acuminatum, contributed above 90 per cent of the seeds to the soil seed bank. On the basis of seed characteristics and seed reduction amount during the growing season, we concluded that it represents a mixture of persistent and transient seeds in different proportions. More seeds accumulated under the canopies than outside the mature shrub cover, but no significant difference was found the younger cover. These results suggest that shrub size and age had an important role in augmenting seed abundance of the soil seed bank but not the species richness. We confirmed the important effects of shrub cover on seed accumulation and vegetation recovery, especially the value of more mature age vegetation for countering land degradation. Copyright © 2012 John Wiley & Sons, Ltd.     

金安桥水电站弃渣场土壤种子库与地面植被的关系

通过研究金安桥水电站废弃渣场种子库和周边群落种子库以及渣场地面和周边群落地面植被的变化特征,结果表明：①周边群落土壤种子库和地面植物的物种丰富度大于渣场,从地面与种子库的物种组成分析得出,渣场植被恢复正处于由一年生草本植物向多年生草本植物过渡的阶段;②周边群落地面植物与土壤种子库的物种丰富度（30,33）及数量（186,1962）,渣场地面植物与土壤种子库的物种丰富度（10,12）及数量（35,1784）;③由于植物种子寿命短,加之土壤环境差,导致周边群落地面植物与种子库相似度（0．09）和渣场地面植物与种子库的相似度（0）差异性大。     

Earthworm-mediated maternal effects on seed germination and seedling growth in three annual plants

Many ecological studies have pointed out maternal effects in plants and shown that plant maternal environment influences germination of their seed and subsequent seedling growth. However, few have tested for maternal effects induced by soil macroorganisms. We tested whether two earthworm species (Aporrectodea caliginosa and Lumbricus terrestris) trigger such maternal effects on seed germination and seedling growth of three plant species (Veronica persica, Poa annua and Cerastium glomeratum). Our results show that, through maternal effects, A. caliginosa enhanced seed germination (V. persica and P. annua) and seedling growth (C. glomeratum and P. annua) while L. terrestris reduced seed germination only in V. persica. In some cases, the increase in germination rates of seeds produced in the presence of earthworms was associated with a reduction of nitrogen content in seeds. These results show that earthworms induce maternal effects in plants and that the size and direction of these effects depend on the combination of plant and earthworm species.     

Soil seed banks in the Arasbaran Protected Area of Iran and their significance for conservation management

The Arasbaran Protected Area is an important but little studied nature reserve in NW Iran. In the past habitat heterogeneity and biodiversity have been threatened by over-exploitation; in the future depopulation may lead to partial abandonment. The soil seed bank was examined to see whether there are sufficient species represented as persistent seeds to reconstitute the vegetation in the event of habitat deterioration. There were differences between seed banks from different soils and vegetation types but essentially all soil seed banks contained few species and low densities of seed. Moreover, the species in the seed bank were unrepresentative of the vegetation. During future periods of changing land use it will be important to maintain the existing vegetation. There are however indications of successful recent conservation management. Annuals are abundant in the woodland seed bank but absent from the vegetation. The forest is recovering from the disturbance of collecting fire-wood.     

Keystone effects of the endangered Stephens' kangaroo rat (Dipodomys stephensi)

Kangaroo rats (Dipodomys) have been argued to exert keystone effects because they interact strongly with other species, and their removal results in major changes in community structure. We evaluated the nature and magnitude of such impacts by the endangered Stephens' kangaroo rat (Dipodomys stephensi), employing an exclosure experiment to determine how the removal of this species impacted relative cover by vegetation, plant species diversity, abundance of a dominant annual plant genus (Erodium), and predation of seeds from artificial trays. Fifteen months of D. stephensi removal resulted in significant increases in herb cover and Erodium abundance, and significant declines in bare ground and in seed predation. These results support the hypothesis that D. stephensi has major impacts on plant community structure and seems to function as a keystone species. It remains to be seen how the loss of this species from much of its range will influence local community composition and dynamics.     

Assessing limitations on population growth in two critically endangered Acacia taxa

Flowering phenology, soil seedbank and the impact of fire, weeds and grazing were investigated in two rare Acacia taxa restricted to the agricultural district north of Perth Western Australia. Acacia aprica is known from six extant populations and one extinct population. Five of the extant populations are restricted to linear road reserves with the sixth population located in a small native vegetation remnant. Acacia cochlocarpa ssp. cochlocarpa is known from a single population also situated on a road verge. Size class structure, levels of canopy death and an absence of juveniles indicated that all populations are in decline. Flowering intensity and success varied between populations and years in A. aprica and between years in A. cochlocarpa ssp. cochlocarpa. Seed bank analysis indicated that seeds were patchily distributed while experimental fires demonstrated that such events could break seed dormancy and promote germination. Both taxa have similar germination physiologies and showed increased germination after seeds were exposed to heat. Following emergence, however, competition with annual weeds had a negative impact on seedling growth and survival in both taxa. While vertebrate grazing had some influence, weeds were the major inhibitory influence on recruitment. Reduced fire frequencies since fragmentation may be responsible for population decline in both taxa but other site specific factors such as weeds and grazing may affect the establishment of seedlings following fire.     

Control of one invasive plant species allows exotic grasses to become dominant in northern Great Plains grasslands

Decline of leafy spurge (Euphorbia esula) in the northern Great Plains of the US is generally viewed as a success story for biological control, but quality of the vegetation that survived the infestation is key to recovery of ecosystem function. In addition, effects of other invasive species, notably cool-season exotic grasses, must be taken into account. Objectives of this study were (1) to evaluate direction and significance of changes in biomass of native and exotic grasses, forbs, and leafy spurge and in plant species composition following control of leafy spurge by flea beetles and (2) to evaluate the relative effects of leafy spurge and exotic grasses on biomass of native grasses, biomass of forbs, and richness of native species. We monitored species composition (1998-2003 and 2008) and biomass (2000, 2002, 2003 and 2008) of these groups on spurge-infested and noninfested permanent plots at three sites with unbroken prairie sod in North Dakota, USA. We found little evidence, in terms of species richness or biomass of native grasses or forbs, that leafy spurge was being replaced by desirable native species, although desirable as well as weedy and exotic species were characteristic of 2008 vegetation at all three sites. Structural equation models revealed that leafy spurge had temporally intermittent negative effects on forb biomass and species richness, but no effects on native grasses. In contrast, exotic grass had consistently strong, negative effects on native grass biomass, as well as stronger negative effects than leafy spurge on native species richness. Although substantial native plant diversity remains at these sites, exotic grasses pose an important threat to these crucial building blocks of native prairie ecosystems.