Seed and establishment limitation contribute to long-term native forb declines in California grasslands

The effects of exotic species invasions on biodiversity vary with spatial scale, and documentation of local-scale changes in biodiversity following invasion is generally lacking. Coupling long-term observations of local community dynamics with experiments to determine the role played by exotic species in recruitment limitation of native species would inform both our understanding of exotic impacts on natives at local scales and regional-scale management efforts to promote native persistence. We used field experimentation to quantify propagule and establishment limitation in a suite of native annual forbs in a California reserve, and compared these findings to species abundance trends within the same sites over the past 48 years. Observations at 11 paired sites (inside and outside the reserve) indicated that exotic annual plants have continued to increase in abundance over the past 48 years. This trend suggests the system has not reached equilibrium > 250 years after exotic species began to spread, and 70 years after livestock grazing ceased within the reserve. Long-term monitoring observations also indicated that six native annual forb species went extinct from more local populations than were colonized. To determine the potential role of exotic species in these native plant declines, we added seed of these species into plots adjacent to monitoring sites where plant litter and live grass competition were removed. Experimental results suggest both propagule and establishment limitation have contributed to local declines observed for these native forbs. Recruitment was highest at sites that had current or historical occurrences of the seeded species, and in plots where litter was removed. Grazing history (i.e., location within or outside the reserve) interacted with exotic competition removal, such that removal of live grass competition increased recruitment in more recently grazed sites. Abundance of forbs was positively related to recruitment, while abundance of exotic forbs was negatively related. Thus, exotic competition is likely only one factor contributing to local declines of native species in invaded ecosystems, with a combination of propagule limitation, site quality, and land use history also playing important and interactive roles in native plant recruitment.     

Seed arrival and ecological filters interact to assemble high-diversity plant communities

Two prominent mechanisms proposed to structure biodiversity are niche-based ecological filtering and chance arrival of propagules from the species pool. Seed arrival is hypothesized to play a particularly strong role in high-diversity plant communities with large potential species pools and many rare species, but few studies have explored how seed arrival and local ecological filters interactively assemble species-rich communities in space and time. We experimentally manipulated seed arrival and multiple ecological filters in high-diversity, herbaceous-dominated groundcover communities in longleaf pine savannas, which contain the highest small-scale species richness in North America (up to > 40 species/m2). We tested three hypotheses: (1) local communities constitute relatively open-membership assemblages, in which increased seed arrival from the species pool strongly increases species richness; (2) ecological filters imposed by local fire intensity and soil moisture influence recruitment and richness of immigrating species; and (3) ecological filters increase similarity in the composition of immigrating species. In a two-year factorial field experiment, we manipulated local fire intensity by increasing pre-fire fuel loads, soil moisture using rain shelters and irrigation, and seed arrival by adding seeds from the local species pool. Seed arrival increased species richness regardless of fire intensity and soil moisture but interacted with both ecological filters to influence community assembly. High-intensity fire decreased richness of resident species, suggesting an important abiotic filter. In contrast, high-intensity fire increased recruitment and richness of immigrating species, presumably by decreasing effects of other ecological filters (competition and resource limitation) in postfire environments. Drought decreased recruitment and richness of immigrating species, whereas wet soil conditions increased recruitment but decreased or had little effect on richness. Moreover, some ecological filters (wet soil conditions and, to a lesser extent, high-intensity fire) increased similarity in the composition of immigrating species, illustrating conditions that influence deterministic community assembly in species-rich communities. Our experiment provides insights into how dispersal-assembly mechanisms may interact with niche-assembly mechanisms in space (spatial variation in disturbance) and time (temporal variation in resource availability) to structure high-diversity communities and can help guide conservation of threatened longleaf pine ecosystems in the face of habitat fragmentation and environmental change.     

Soil vs. canopy seed storage and plant species coexistence in species-rich Australian shrublands

The fire-prone shrublands of southwestern Australia are renowned for their high plant species diversity and prominence of canopy seed storage (serotiny). We compared species richness, abundance, and life history attributes for soil and canopy seed banks in relation to extant vegetation among four sites with different substrate conditions and high species turnover (50-80%) to identify whether this unusual community-level organization of seed storage might contribute to maintenance of high species richness. Soil seed bank (SSB) densities were low to moderate (233-1435 seeds/m2) compared with densities for other Mediterranean-type vegetation and were lowest for sites with highest canopy seed bank (CSB) species richness and lowest nutrient availability, but not richness or abundance of resprouters. Annuals were infrequent in the lowest nutrient sites, but there was no evidence that small SSB size was due to low seed inputs or a trade-off between seed production/storage and seed size in response to low nutrient availability. Sorensen's similarity between SSB and extant vegetation was 26-43% but increased to 54-57% when the CSB was included, representing levels higher than reported for most other ecosystems. Resprouting species were well represented in both the SSB and CSB, and there was no evidence for lower seed production in resprouters than in non-sprouters overall. The SSB and CSB held no species in common and were characterized by markedly different seed dispersal attributes, with winged or small seeds in the CSB and seeds dispersed by ants, birds, and wind (though none with wings) in the SSB. There was no evidence of spatial differentiation in the distribution of seeds of SSB species between vegetated and open microsites that might facilitate species coexistence, but most woody non-sprouters showed aggregation at scales of 1-2 m, implying limited seed dispersal. High similarity between overall seed bank (SSB + CSB) and extant species composition, high number of resprouting species, and seed dispersal processes before (SSB) and after fire (CSB) leading to differential spatial aggregation of post-fire recruits from the two seed bank types may buffer species composition against rapid change and provide a mechanism for maintaining species coexistence at the local scale.     

Modifying native and exotic species richness correlations: the influence of fire and seed addition.

An important goal in restoration is to increase the richness of native species while reducing exotic species. However, native species richness is often positively correlated with exotic species richness. In a grassland-savanna system in Michigan (USA), we show that management that focuses on changing the nature of the exotic-native richness relationship can be used to restore native communities. Native and exotic species richnesses were positively correlated, likely due to a shared coupling with aboveground live biomass (a surrogate for productivity). The addition of native seed shifted the exotic-native richness relationship from a linear positive to a monotonic relationship: in areas of intermediate levels of exotic species richness, seed addition increased native diversity without an associated effect on exotic diversity, but in areas of high or low exotic richness, it did not affect native species richness. Prescribed burning broke the correlation between native and exotic richness with no consistent effect on the richness of either group. However, when burning was combined with native-seed addition, the relationship between native and exotic richness was maintained and was shifted upwards, enhancing native recruitment. Although aboveground productivity was strongly related to species richness across the landscape, changes in productivity did not drive these shifts.     

Recovery of plant diversity following N cessation: effects of recruitment, litter, and elevated N cycling

Plant species richness has declined and composition shifted in response to elevated atmospheric deposition of biologically active nitrogen over much of the industrialized world. Litter thickness, litter nitrogen (N) content, and soil N mineralization rates often remain elevated long after inputs cease, clouding the prospects that plant community diversity and composition would recover should N inputs be reduced. Here we determined how N cycling, litter accumulation, and recruitment limitation influenced community recovery following cessation of long-term N inputs to prairie-like grasslands. We alleviated each of these potential inhibitors through a two-year full-factorial experiment involving organic carbon addition, litter removal, and seed addition. Seed addition had the largest effect on increasing seedling and species numbers and may be necessary to overcome long-term burial of seeds of target perennial grassland species. Litter removal increased light availability and bare sites for colonization, though it had little effect on reducing the biomass of competing neighbors or altering extractable soil N. Nonetheless, these positive influences were enough to lead to small increases in species richness within one year. We found that, although C addition quickly altered many factors assumed favorable for the target community (decreased N availability and biomass of nearby competitors, increased light and site availability), these changes were insufficient to positively impact species richness or seedling numbers over the experimental duration. However, only carbon addition had species-specific effects on the existing plant community, suggesting that its apparent limited utility may be more a result of slow recovery under ambient recruitment rather than from a lack of a restorative effect. There were dramatic interactions among treatments, with the positive effects of litter removal largely negated by carbon addition, and the positive effects of seed addition generally amplified by litter removal. It remains unclear whether each mechanism explored here will induce community recovery, but over different temporal scales. Long-term monitoring will help resolve these remaining questions. Regardless, our results suggest that reversal of species loss and compositional shifts from N deposition in prairies may be more inhibited by habitat fragmentation, recruitment limitation, and long-term suppression of fire than from continued effects of elevated N.     

Dispersal, competition, and shifting patterns of diversity in a degraded oak savanna

Diversity is a balance between processes that add and limit species (e.g., dispersal vs. competition), but reconciling their contributions remains a challenge. Recruit-ment-based models predict that dispersal barriers are most limiting for diversity, while competition-based models predict that dispersal matters only when competition is minimized. Testing these models is difficult because their influence varies with scale and site productivity. In a degraded oak savanna, we used plot-level (seed additions, burning) and site-level (proportions of regional functional groups found locally) analyses in areas with variable soil depth to examine how dispersal and competition influence diversity. At the plot level, added species persisted where they were formerly absent, but few established naturally despite fire-induced resource enrichment and nearby populations, revealing the importance of dispersal limitation for diversity. This result did not vary with soil depth or standing crop. Although competition could not prevent establishment in unburned plots, it significantly lowered survival, indicating that resource limitations exacerbate dispersal inefficiencies. At the site level, the concordance between regional and local diversity for native species was associated with soil depth heterogeneity, not dispersal or competition. This suggests that persistence is determined primarily by the influence of the environment on population demographics. Given that the formation of new populations is unlikely, those remaining appear to be confined to optimal habitat where they resist competitive or stochastic displacement, possibly explaining why species loss is rare despite substantial habitat loss and invasion. For exotics, there was no relationship between diversity and soil depth heterogeneity. Annuals with presumed dispersal capabilities were significantly overrepresented in all sites while perennial forbs, the largest regional functional group, were significantly underrepresented. We interpret the native-exotic discrepancies as reflecting the recent arrival of exotics (150 years ago), suggesting that local establishment occurs slowly even for species with regional prevalence. The accumulation lag may be explained by the need for founder populations to be demographically stable; otherwise persistence requires continual immigration favoring overrepresentation by dispersers. Our findings support the view that dispersal limitation restricts diversity within plant communities, but suggests that the impacts of environment on demographic performance ultimately determine the pattern and rate of community assembly.     

Bridging the generation gap in plants: pollination, parental fecundity, and offspring demography

Despite extensive study of pollination and plant reproduction on the one hand, and of plant demography on the other, we know remarkably little about links between seed production in successive generations, and hence about long-term population consequences of variation in pollination success. We bridged this "generation gap" in Ipomopsis aggregata, a long-lived semelparous wildflower that is pollinator limited, by adding varying densities of seeds to natural populations and following resulting plants through their entire life histories. To determine whether pollen limitation of seed production constrains rate of population growth in this species, we sowed seeds into replicated plots at a density that mimics typical pollination success and spacing of flowering plants in nature, and at twice that density to mimic full pollination. Per capita offspring survival, flower production, and contribution to population increase (lambda) did not decline with sowing density in this experiment, suggesting that typical I. aggregata populations freed from pollen limitation will grow over the short term. In a second experiment we addressed whether density dependence would eventually erase the growth benefits of full pollination, by sowing a 10-fold range of seed densities that falls within extremes estimated for the natural "seed rain" that reaches the soil surface. Per capita survival to flowering and age at flowering were again unaffected by sowing density, but offspring size, per capita flower production, and lambda declined with density. Such density dependence complicates efforts to predict population dynamics over the longer term, because it changes components of the life history (in this case fecundity) as a population grows. A complete understanding of how constraints on seed production affect long-term population growth will hinge on following offspring fates at least through flowering of the first offspring generation, and doing so for a realistic range of population densities.     

Herbivory limits recruitment in an old-field seed addition experiment

Environmental variability can promote coexistence by creating establishment sites for rare plants, but low diversity in anthropogenic grasslands suggests that this variability may be eliminated (homogenization hypothesis) or inaccessible (barrier hypothesis). We explore these alternatives on the northern Great Plains, where 11 million hectares have been transformed by multiple environmental changes, but the causes of species loss are unclear. In a degraded grassland, we increased environmental variability by manipulating competition and herbivory along gradients of fertility and disturbance, and we circumvented dispersal barriers by adding 1.2 million seeds of five functionally distinct species at varying densities. The experiment ended after 12 weeks due to the direct and indirect effects of unapparent small native herbivores, which were barriers to population establishment by the added species. The direct cause of recruitment failure was browsing. The indirect cause was associated with competition from invasive plants that appeared to be more tolerant or resistant to herbivory. Variability in fertility, disturbance, propagule pressure, and competition had relatively minor impacts on colonization by the added species because herbivores controlled recruitment in most environments. Recruitment outside the herbivore exclosures was mostly by unpalatable exotics, suggesting a possible link between invasion success and herbivore resistance for some introduced plants.     

Diversity, host affinity, and distribution of seed-infecting fungi: a case study with Cecropia

Recruitment limitation has been proposed as an important mechanism contributing to the maintenance of tropical tree diversity. For pioneer species, infection by fungi significantly reduces seed survival in soil, potentially influencing both recruitment success and adult distributions. We examined fresh seeds of four sympatric Cecropia species for evidence of fungal infection, buried seeds for five months in common gardens below four C. insignis crowns in central Panama, and measured seed survival and fungal infection of inviable seeds. Seed survival varied significantly among species and burial sites, and with regard to local (Panama) vs. foreign (Costa Rica) maternal seed sources. Fresh seeds contained few cultivable fungi, but > 80% of soil-incubated seeds were infected by diverse Ascomycota, including putative pathogens, saprophytes, and endophytes. From 220 isolates sequenced for the nuclear internal transcribed spacer region (ITS), 26 of 73 unique genotypes were encountered more than once. Based on the most common genotypes, fungal communities demonstrate host affinity and are structured at the scale of individual crowns. Similarity among fungal communities beneath a given crown was significantly greater than similarity among isolates found under different crowns. However, the frequency of rare species suggests high fungal diversity and fine-scale spatial heterogeneity. These results reveal complex plant-fungal interactions in soil and provide a first indication of how seed survival in tropical forests may be affected by fungal community composition.     

Invasive competitor and native seed predators contribute to rarity of the narrow endemic Astragalus sinuatus Piper

The conservation of rare plant species hinges on our ability to identify the underlying mechanisms that limit rare plant populations. Theory on rarity suggests that both predispersal seed predation and competition can be important mechanisms influencing abundance and/or distribution of rare plant populations. Yet few studies have tested these interactions, and those that have evaluated each mechanism independently. Astragalus sinuatus Piper (Whited's milkvetch) is a narrow endemic plant species restricted to eight populations within a 10-km2 area in eastern Washington. We used experimental and observational methods to test the effects of native insect predispersal seed predators and an invasive grass (Bromus tectorum L. [cheatgrass]) on seed set and population density of A. sinuatus. We quantified per capita seed production and pod predation rates across four sites and among four years. Seed predation rates were high across four sites (66-82%) and all years (65-82%). Experimental reduction of predispersal seed predators significantly increased per capita seed set of A. sinuatus (164-345%) at two experimental sites. Concurrently, two seed addition experiments demonstrated the effect of seed loss and presence of B. tectorum on seedling recruitment and establishment of A. sinuatus over four growing seasons. In the first seed addition experiment, we found no difference in recruitment and establishment between low (40) and high (120) seed addition levels. In the second addition experiment (one level of addition; 40 seeds), we found that recruitment and survivorship increased 200% in plots where B. tectorum was removed compared to plots where B. tectorum was present. Thus, seed addition had no impact in the presence of B. tectorum; conversely, in the absence of B. tectorum, seed addition was highly effective at increasing population numbers. Results suggest that, in areas where B. tectorum is present, recruitment is site limited, and it is seed limited when B. tectorum is absent. We recommend that managers reduce B. tectorum in an effort to increase population growth of A. sinuatus; in areas where B. tectorum is absent, short-term reduction of insect predators should be considered as a strategy to increase population growth of this rare species.     

全尺度人工湿地中植物多样性对生产力与多样性效应的影响

在中国东南部的全尺度复合垂直流人工湿地中开展2年的植物多样性实验,以研究植物多样性（包括植物物种丰富度和植物组成）对群落生产力与多样性效应（即互补效应、选择效应和净多样性效应）的影响及其产生机制。结果表明,2007年物种丰富度与群落生产力呈线形正相关,而2008年显著的单峰格局,其关系式为：y=-0.213x2＋3.455x＋15.192（R=0.215）。2008年物种丰富度与互补效应呈显著地线形负相关,而2007年呈单峰格局,其关系式为：y=-0.389x2＋6.974x-10.707（R=0.247）,而且2007年与2008年的互补效应与生产力都呈显著的正相关,表明互补效应对生产力的提高有重要作用。然而,2007年与2008年物种丰富度与选择效应之间均没有显著相关性,且选择效应与群落生产力之间也没有显著相关性,表明选择效应对生产力的提高作用不显著。2007年与2008年中物种组成对生产力、互补效应、选择效应与净多样性效应均有显著影响,说明人工湿地的植物配置对其生态系统功能的维持尤为重要。2008年物种丰富度与净多样性效应呈极显著地线形负相关,而2007年呈显著单峰格局,其关系式为：y=-0.329 x2＋5.968 x-12.659（R=0.234）,这种趋势主要是由于植物多样性-生态系统功能关系的影响因素（如物种的竞争力和生态位）在2年中有所变化。同时,2007年与2008年的多样性净效应与生产力都呈显著正相关关系,表明生产力与多样性净效应的变化趋势是同步的。与抽样效应假说不同的是,本实验中单种最高产物种（芦竹）在混种时没有表现出高产,主要是由于生长的分配、资源的竞争力与环境的变化等。     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Abstract: Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed‐deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird‐dispersed, fleshy‐fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit‐rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird‐dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed‐to‐seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy‐fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed‐survival expectancies at microhabitats to preserve plant‐population dynamics and community structure in fragmented landscapes.     

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.     

Hunting Increases Dispersal Limitation in the Tree Carapa procera, a Nontimber Forest Product

Abstract:  The sustainability of seed extraction from natural populations has been questioned recently. Increased recruitment failure under intense seed harvesting suggests that seed extraction intensifies source limitation. Nevertheless, areas where more seeds are collected tend to also have more intense hunting of seed-dispersing animals. We studied whether such hunting, by limiting disperser activity, could cause quantitative dispersal limitation, especially for large crops and for crops in years of high seed abundance. In each of four Carapa procera (Meliaceae) populations in French Guiana and Surinam, two with hunting and two without, we compared seed fate for individual trees varying in crop size in years of high and low population-level seed abundance. Carapa seeds are a nontimber forest product and depend on dispersal by scatter-hoarding rodents for survival and seedling establishment. Hunting negatively affected the proportion of seeds dispersed and caused greater numbers of seeds to germinate or be infested by moths below parent trees, where they would likely die. Hunting of seed-dispersing animals disproportionally affected large seed crops, but we found no additional effect of population-level seed abundance on dispersal rates. Consistently lower rates of seed dispersal, especially for large seed crops, may translate to lower levels of seedling recruitment under hunting. Our results therefore suggest that the subsistence hunting that usually accompanies seed collection is at the cost of seed dispersal and may contribute to recruitment failure of these nontimber forest products. Seed extraction from natural populations may affect seedling recruitment less if accompanied by measures adequately incorporating and protecting seed dispersers.     

Seed predation and selection exerted by a seed predator influence subalpine tree densities

Strongly interacting species often have pronounced direct and indirect effects on other species. Here we focus of the effects of pine squirrels (Tamiasciurus spp.), which are a dominant pre-dispersal seed predator of many conifers including limber pines (Pinus flexilis) and whitebark pines (P. albicaulis). Pine squirrels depress seed abundance by harvesting most limber and whitebark pine cones on their territories. Pine squirrels further reduce seed availability for Clark's Nutcrackers (Nucifraga columbiana), the primary seed disperser of these pines, because selection exerted by pine squirrels has reduced the number of seeds per cone and causes seeds to be less accessible. We predicted that, if fewer seeds were available for dispersal by nutcrackers, pine recruitment should be suppressed in areas with pine squirrels. In support of this prediction, stand densities were about two times greater in areas where pine squirrels are absent than in areas where they are present. Alternative explanations that we considered do not account for these differences; however, precipitation may limit stand densities in the absence of seed limitation by pine squirrels. In sum, pine squirrels apparently depress limber and whitebark pine stand densities, with the potential for ecosystem impacts because these pines are foundation species within Western subalpine ecosystems.     

High richness and dense seeding enhance grassland restoration establishment but have little effect on drought response

Restorations commonly utilize seed addition to formerly arable lands where the development of native plant communities is severely dispersal limited. However, variation in seed addition practices may profoundly affect restoration outcomes. Theory and observations predict that species-rich seed mixtures and seeding at high densities should enhance native plant community establishment, minimize exotic species cover, and may promote resistance and resilience to, and recovery from, environmental perturbations. We studied the post-seeding establishment of native plant communities in large grassland restoration plots, which were sown at two densities crossed with two levels of species richness on formerly arable land in Nebraska, USA, and their responses to drought. To evaluate drought resistance, recovery, and resilience of restored plant communities, we erected rainfall manipulation structures and tracked the response of seeded species cover and total plant biomass during experimental drought relative to controls and in the post-drought growing season. High seed richness and high-density seeding treatments resulted in greater richness and cover of native, seeded species per 0.5 m2 compared to low-richness and low-density treatments. Cover differences in response to seed mixture richness were driven by native forbs. Richness and cover of exotic species were lowest in high-richness and high-density treatments. We found little evidence of differential drought resistance, recovery, and resilience among seeding treatments. Increases in exotic species across years were restricted to drought subplots, and were not affected by seeding treatments. Grassland restoration was generally enhanced and exotic cover reduced both by the use of high-richness seed mixtures and high-density seeding. Given the lack of restoration treatment effects on the resistance, recovery, or resilience of seeded species exposed to drought, and the increases in exotic species following drought, other forms of active management may be needed to produce restored plant communities that are robust to climate change.     

Effects of productivity on biodiversity in forest ecosystems across the United States and China

In the global campaign against biodiversity loss in forest ecosystems, land managers need to know the status of forest biodiversity, but practical guidelines for conserving biodiversity in forest management are lacking. A major obstacle is the incomplete understanding of the relationship between site primary productivity and plant diversity, due to insufficient ecosystem‐wide data, especially for taxonomically and structurally diverse forest ecosystems. We investigated the effects of site productivity (the site's inherent capacity to grow timber) on tree species richness across 19 types of forest ecosystems in North America and China through 3 ground‐sourced forest inventory data sets (U.S. Forest Inventory and Analysis, Cooperative Alaska Forest Inventory, and Chinese Forest Management Planning Inventory). All forest types conformed to a consistent and highly significant (P < 0.001) hump‐shaped unimodal relationship, of which the generalized coefficients of determination averaged 20.5% over all the forest types. That is, tree species richness first increased as productivity increased at a progressively slower rate, and, after reaching a maximum, richness started to decline. Our consistent findings suggest that forests of high productivity would sustain few species because they consist mostly of flat homogeneous areas lacking an environmental gradient along which a diversity of species with different habitats can coexist. The consistency of the productivity–biodiversity relationship among the 3 data sets we examined makes it possible to quantify the expected tree species richness that a forest stand is capable of sustaining, and a comparison between the actual species richness and the sustainable values can be useful in prioritizing conservation efforts.     

部分菊科入侵种种子(瘦果)的萌发能力和幼苗建群特性

对入侵种种子生物学各方面的了解有助于更好地预测入侵趋势和评估可持续管理的策略.研究通过对部分菊科入侵种种子(瘦果)进行的室内萌发实验和土壤埋藏实验,通过与相关研究的对比和分析,对这些入侵种的萌发能力和幼苗建群特性进行了研究.10个入侵种种子室内萌发实验的结果表明,在25℃、12h光照的情况下,除钻形紫菀和一年蓬以外,被研究的其它8个种的终萌发率超过了50%,而且都显示出早期快速萌发的特点.用SPSS的曲线拟合工具对10个种的累积萌发趋势进行了曲线回归分析,对应的最佳模型均为三次曲线方程.不同土壤埋藏深度下幼苗出苗实验的结果表明,菊科8个入侵种的种子在表土的出苗率均最高,随着覆土厚度的增加,种子的出苗率逐渐降低,当覆土厚度超过3 cm以上时,所有种子均不能出苗.种子在不同的埋藏深度的出苗情况与种子的大小有关,较大种子破土能力相对较强;种子在不同的埋藏深度的出苗情况与种子本身的萌发能力也相关.土壤埋藏期限的实验结果表明,随埋藏时期的增加,小蓬草、苏门白酒草和一年蓬3个人侵种种子的活力逐步降低,在土壤中埋藏12个月后,其活力分别为21.34%、18.15%和27.38%.这些入侵种较广的种子萌发温度适应范围和高萌发率为防除工作带来了困难.在生产上,可通过不同时期的翻耕、将种子深埋等措施减少这些入侵种的危害.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.