Biocontrol attack increases pollen limitation under some circumstances in the invasive plant Centaurea solstitialis

Herbivore damage often deters pollinator visitation and many invasive plants in North America are pollinator-dependent. This has important implications for the biological control of invasive plants because it means that agents that deter pollinators may have a larger than expected impact on the plant. Yet interactions between pollinators and biocontrol agents are rarely evaluated. Centaurea solstitialis, one of the most problematic invasive species in California, is dependent on pollinators for reproduction. I factorially manipulated infection by a biocontrol pathogen and pollen supplementation to test for (1) pollen limitation in C. solstitialis, (2) whether infection increased pollen limitation, and (3) whether this varied across a soil moisture gradient. Plants growing on north-facing slopes where soil moisture was higher experienced mild pollen limitation in the absence of the pathogen and more pronounced pollen limitation when they were infected. Plants on drier south-facing slopes did not suffer from pollen limitation but instead appeared to suffer from resource limitation. Pathogen infection directly reduced seed set in C. solstitialis by 67–72 %. On north-facing slopes, infection had an additional, indirect effect by increasing the degree of pollen limitation plants experienced. The trait that mediates this indirect pathogen–pollinator interaction is the number of inflorescences plants produced: infected plants made fewer inflorescences which led to greater pollen limitation. Although in the present study this outcome is dependent on abiotic factors that vary over small spatial scales, exploiting other invasive plants’ dependence on pollinators by selecting agents that deter visitation may enhance agent impact.     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

Consumer pressure, seed versus safe-site limitation, and plant population dynamics

Plants often suffer reductions in fecundity due to insect herbivory. Whether this loss of seeds has population-level consequences is much debated and often unknown. For many plants, particularly those with long-lived seedbanks, it is frequently asserted that herbivores have minimal impacts on plant abundance because safe-site availability rather than absolute seed number determines the magnitude of future plant recruitment and hence population abundance. However, empirical tests of this assertion are generally lacking and the interplay between herbivory, spatio-temporal variability in seed- or safe-site-limited recruitment, and seedbank dynamics is likely to be complex. Here we use a stochastic simulation model to explore how changes in the spatial and temporal frequency of seed-limited recruitment, the strength of density-dependent seedling survival, and longevity of seeds in the soil influence the population response to herbivory. Model output reveals several surprising results. First, given a seedbank, herbivores can have substantial effects on mean population abundance even if recruitment is primarily safe-site-limited in either time or space. Second, increasing seedbank longevity increases the population effects of herbivory, because annual reductions in seed input due to herbivory are accumulated in the seedbank. Third, population impacts of herbivory are robust even in the face of moderately strong density-dependent seedling mortality. These results imply that the conditions under which herbivores influence plant population dynamics may be more widespread than heretofore expected. Experiments are now needed to test these predictions. Received: 3 November 1999 / Accepted: 15 February 2000     

Comparison of the effect of elevated CO2 on an invasive species (Centaurea solstitialis) in monoculture and community settings

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) is likely to change the species composition of plant communities. To investigate whether growth of a highly invasive plant species, Centaurea solstitialis (yellow starthistle), was affected by elevated [CO₂], and whether the success of this species would increase under CO₂ enrichment, I grew the species in serpentine soil microcosms, both as a monoculture and as a component of a grassland community. Centaurea grown in monoculture responded strongly to [CO₂] enrichment of 350 mol mol^–1, increasing aboveground biomass production by 70%, inflorescence production by 74%, and midday photosynthesis by an average of 132%. When grown in competition with common serpentine grassland species, Centaurea responded to CO₂ enrichment with similar but nonsignificant increases (+69% aboveground biomass, +71% inflorescence production), while total aboveground biomass of the polyculture increased by 28%. Centaurea's positive CO₂ response in monoculture and parallel (but non-significant) response in polyculture provoke questions about possible consequences of increasing CO₂ for more typical California grasslands, where the invader already causes major problems.     

Spatial variation in seed limitation of plant species richness and population sizes in floodplain tallgrass prairie

The relative importance of seed availability versus biotic interactions that affect early life stages in limiting plant population sizes and determining composition of plant communities is a central debate in plant ecology. We conducted a seed addition experiment in restored tallgrass prairie in central Kansas to determine (1) whether addition of seed of 18 native forb species produced persistent (three growing seasons) increases in the species' population sizes and plant species richness, (2) what properties of recipient communities best explained spatial variation in added species' establishment, and (3) whether seed size explained interspecific patterns in establishment success. Adding seed led to persistent increases in the number of added species present and in plant species richness at one of three sites. Increased species richness at the one site where community composition was structured by seed availability largely resulted from greater densities of four species. Seed size did not predict species' establishment success. Pre-existing plant species richness was correlated with added species' establishment success, but the direction of the relationship (positive vs. negative) varied among sites. Living aboveground plant biomass in experimental plots in the year of seed addition was negatively correlated with the number of added species established three years later. Our results provide further evidence for large spatial variation in seed limitation of plant community composition. Surprisingly, mean light availability and heterogeneity in light, both important parameters in conceptual models of grassland plant coexistence, did not predict the response of the recipient plant community to seed addition as well as pre-existing plant species richness and living aboveground biomass.     

Functional traits but not environmental gradients explain seed weight in Mongolian plant species

• Seed weight varies by several orders of magnitude among vascular plant species. However, the importance of potential drivers such as environmental conditions and plant functional traits have rarely been assessed for a larger taxonomic sample.
• We collected seeds of 148 species from 237 sites spread across Mongolia and compared their weight among the major zonal vegetation types, taxonomic groups and a set of functional traits (growth form, dispersal mode, fruit type, storage organs and palatability).
• Seed weight strongly varied among all functional traits and taxonomic groups, but no differences among vegetation zones were detected.
• These results suggest a low impact of environmental conditions on the evolution of seed weight, contrasting the strong phylogenetic signal.

     

The invasive plant species Centaurea maculosa alters arbuscular mycorrhizal fungal communities in the field

While several recent studies have described changes in microbial communities associated with exotic plant invasion, how arbuscular mycorrhizal fungi (AMF) communities respond to exotic plant invasion is not well known, despite the salient role of this group in plant interactions. Here, we use molecular methods (terminal restriction fragment length polymorphism analyses based on the large subunit of the rRNA gene) to examine AMF community structure in sites dominated by the invasive mycorrhizal forb, Centaurea maculosa Lam. (spotted knapweed), and in adjacent native grassland sites. Our results indicate that significant AMF community alteration occurs following C. maculosa invasion. Moreover, a significant reduction in the number of restriction fragment sizes was found for samples collected in C. maculosa-dominated areas, suggesting reduced AMF diversity. Extraradical hyphal lengths exhibited a significant, on average 24%, reduction in C. maculosa-versus native grass-dominated sites. As both AMF community composition and abundance were altered by C.maculosa invasion, these data are strongly suggestive of potential impacts on AMF-mediated ecosystem processes. Given that the composition of AMF communities has the potential to differentially influence different plant species, our results may have important implications for site restoration after weed invasion.     

Increased seed consumption by biological control weevil tempers positive CO2 effect on invasive plant (Centaurea diffusa) fitness

Predicted increases in atmospheric CO₂ and temperature may benefit some invasive plants, increasing the need for effective invasive plant management. Biological control can be an effective means of managing invasive plants, but the anticipated range in responses of plant–insect interactions to climate change make it difficult to predict how effective biological control will be in the future. Field experiments that manipulate climate within biological control systems could improve predictive power, but are challenging to implement and therefore rare to date. Here, we show that free air CO₂ enrichment in the field increased the fitness of Centaurea diffusa Lam., a problematic invader in much of the western United States. However, CO₂ enrichment also increased the impact of the biological control agent Larinus minutus (Coleoptera: Curculionidae) on C. diffusa fitness. C. diffusa plants flowered earlier and seed heads developed faster with both elevated CO₂ and increased temperature. Natural dispersal of L. minutus into the experimental plots provided a unique opportunity to examine weevil preference for and effects on C. diffusa grown under the different climate change treatments. Elevated CO₂ increased both the proportion of seed heads infested by L. minutus and, correspondingly, the amount of seed removed by weevils. Warming had no detectable effect on weevil utilization of plants. Higher weevil densities on elevated CO₂ plants reduced, but did not eliminate, the positive effects of CO₂ on C. diffusa fitness. Correlations between plant development time and weevil infestation suggest that climate change increased weevil infestation by hastening plant phenology. Phenological mismatches are anticipated with climate change in many plant–insect systems, but in the case of L. minutus and C. diffusa in mixed-grass prairie, a better phenological match may make the biological control agent more effective as CO₂ levels rise.     

Increased population growth rate in invasive polyploid Centaurea stoebe in a common garden

Biological invasions are inherently demographic processes, but trait differences between native and introduced genotypes are rarely linked to population growth rates. Native European Centaurea stoebe occurs as two cytotypes with different life histories (monocarpic diploids, polycarpic tetraploids); however, only tetraploids have been found in its introduced range in North America. In a common garden experiment using artificial populations, we compared the demographic performance of the three geo-cytotypes in the presence and absence of a specialist herbivore using periodic matrix models. We found no difference in population growth rate between the two European cytotypes and no significant effects of herbivory in all geo-cytotypes. However, there was a pronounced increase in population growth rate for North American compared with European tetraploids due to increased seed production and juvenile establishment. These results suggest that genetic drift or rapid evolution, rather than pre-adaptation through polyploidy may explain the invasion success of tetraploids.     

Shoot herbivory on the invasive plant, Centaurea maculosa, does not reduce its competitive effects on conspecifics and natives

Herbivory can have negative, positive, or no effect on plants. However, insect biological control assumes that herbivory will negatively affect the weed and release natives from competition. Centaurea maculosa, an invader in North America, is tolerant to herbivory, and under some conditions, herbivory may increase its competitive effects on natives. Therefore, we investigated two hypotheses: 1) herbivory stimulates compensatory growth by C. maculosa, which increases its competitive effects, and 2) herbivory stimulates the allelopathic effect of C. maculosa. In the greenhouse, Trichoplusia ni shoot herbivory reduced C. maculosa biomass when shoot damage exceeded 40% of the total original leaf area. Conspecific neighbors had no effect on C. maculosa biomass, and the presence of the natives Festuca idahoensis and F. scabrella had a positive effect on C. maculosa. Neighbors did not alter the effects of shoot herbivory. More importantly, even intense shoot herbivory on C. maculosa did not benefit neighboring plants. In a field experiment, clipping 50% of C. maculosa aboveground biomass in the early summer and again in the late summer reduced final biomass by 40% at the end of the season; however, this clipping did not affect total biomass production or reproductive output. Festuca idahoensis neighbors did not increase the effects of clipping, and aboveground damage to C. maculosa did not release F. idahoensis from competition. In the greenhouse we used activated carbon to adsorb allelochemicals, which reduced the competitive effects of C. maculosa on F. idahoensis but not on F. scabrella or other C. maculosa. However, we found no increase in the allelopathic effects of C. maculosa after shoot herbivory. In summary, our results correspond with others indicating that exceptionally high intensities of herbivory are required to suppress C. maculosa growth and reproduction; however, even intense herbivory on C. maculosa does not insure that native bunchgrasses will benefit.     

Reducing dispersal limitation via seed addition increases species richness but not above‐ground biomass

Seed dispersal limitation, which can be exacerbated by a number of anthropogenic causes, can result in local communities having fewer species than they might potentially support, representing a potential diversity deficit. The link between processes that shape natural variation in diversity, such as dispersal limitation, and the consequent effects on productivity is less well known. Here, we synthesised data from 12 seed addition experiments in grassland communities to examine the influence of reducing seed dispersal limitation (from 1 to 60 species added across experiments) on species richness and productivity. For every 10 species of seed added, we found that species richness increased by about two species. However, the increase in species richness by overcoming seed limitation did not lead to a concomitant increase in above‐ground biomass production. This highlights the need to consider the relationship between biodiversity and ecosystem functioning in a pluralistic way that considers both the processes that shape diversity and productivity simultaneously in naturally assembled communities.     

The small‐scale spatiotemporal pattern of the seedbank and vegetation of a highly invasive weed,Centaurea solstitialis: strength in numbers

The dynamics of invasive plant populations are intriguing and informative of the importance of population and community‐level processes. A dominant approach to understanding and describing invasion has been the development of unique hypotheses to explain invasion. However, here we directly explore the relevance of the small‐scale, spatiotemporal pattern in seedbanks and plants of the highly invasive weed, Centaurea solstitialis, to determine whether pattern can be used to contrast predictions associated with the simple ecological hypotheses of seed versus microsite limitations. At three invaded grasslands in California, highly invaded (> 20 adult plants present), invaded (< 10 adults), and uninvaded (no C. solstitialis plants) sites were selected. The spatial pattern of the seedbank was assessed using fine‐scale, 2 cm diameter contiguous cores and geostatistical statistics, and the number of C. solstitialis seeds in the seedbank was recorded in addition to the total community seedbank density. Three of the four critical predictions associated with the seed limitation hypothesis were clearly supported as an explanation for the patterns of C. solstitialis invasion observed in the field. The density of C. solstitialis seeds decreased from high to low extents of invasion, there was no relationship between the community seedbank and C. solstitialis seeds, and the distances between C. solstitialis plants was inversely related to the density of C. solstitialis seeds. However, both the persistent and transient seedbanks of C. solstitialis were spatially aggregated with autocorrelation up to 12 cm² which suggests that aggregation is a consistent attribute of this species in the seedbank regardless of extent of invasion. This basic pattern‐based approach clearly detected an ecological signal of invasive seedbank dynamics and is thus a useful tool for subsequent studies of invasions in grasslands.     

Nitrogen addition affects plant biomass allocation but not allometric relationships among different organs across the globe

全球尺度上氮添加影响植物生物量分配但不影响不同器官间的异速生长关系 生物量在不同器官间的分配是植物对环境变化响应的一个关键生态生理学过程。然而,在全球尺度上有关不同陆地生态系统植物生物量分配对氮沉降响应的认识还比较欠缺。本文通过整合分析333篇已发表文章的5474个观测值,基于“最优分配假说”和“异速分配假说”,评估了全球尺度上氮添加对植物生物量及其在不同器官间分配的影响。结果表明：(1)氮添加显著增加了整株植物或不同器官的生物量,降低了根冠比和根质量分数,但对叶质量分数和茎质量分数无显著影响;(2)氮添加对不同器官质量分数的影响受实验条件、植物功能性状、纬度、氮添加率等因子单独或交互作用的调控;(3)氮添加对生物量在不同器官间的异速分配率无显著影响,表明氮添加导致的根冠比和根质量分数减少是在异速分配模式下由整株植物生物量增加而致。虽然氮添加改变了地上和地下部分的质量分数,但不同器官生物量间异速分配模式的稳定性表明“异速分配假说”能更好地描述植物生物量分配对氮添加的响应规律。该研究结果将有助于深入认识氮沉降环境下植物生物量的分配规律,同时表明将生物量分配纳入有关氮沉降对陆地生态系统影响的预测模型中的重要性。     

Livestock settlement affects shrub abundance via plant functional diversity but not species richness in arid environments

Plant Ecology - Recent theoretical and empirical studies have assumed that livestock grazing acts as a biotic filter favouring plant species with functional traits that confer resistance to...     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

Pollen viability and limitation of seed production in a population of the circumpolar cushion plant, Silene acaulis (Caryophyllaceae)

Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0–53% in hermaphrodite and 0–54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) hand-pollination without removal of other flowers, and (III) open pollination without removal of flowers. Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of 5. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.     

Reproductive assurance through self-fertilization does not vary with population size in the alien invasive plant Datura stramonium

Autonomous self-fertilization is suggested to be associated with invasiveness in plants because it offers reproductive assurance when there is a shortage of suitable mates or pollinators. Given that shortages of mates and pollinators are a common cause of Allee effects in small plant populations, we predict that the benefits of self-fertilization in terms of reproductive assurance should be greatest in small populations. We tested this idea for the invasive herb Datura stramonium , a self-fertilizing species which is also cross-pollinated to some extent by insects (mainly hawkmoths and honeybees). During two consecutive years, we studied 20 and 55 populations, respectively, of different sizes. Untreated flowers of D. stramonium showed high levels of fruit and seed set in all populations studied. Although, fruit and seed set were generally reduced by about 90% in flowers in which self-fertilization was prevented through emasculation, this effect did not vary according to population size. By using a natural color (anthocyanin) dimorphism in 12 populations, we showed that the average outcrossing rate was low (1.3%) and that there was no relationship between outcrossing rate and population size. Pollen removal from flowers also did not vary according to population size, suggesting that the pollinator visitation rate is not lower in small populations. However, decreasing deviations of observed from expected fruit set with population size imply that small populations may have an increased chance of extinction due to demographic stochasticity. Overall, our results suggest that reproductive assurance through self-fertilization in invasive plants may be important for all stages of population establishment, and not just in the founder population.