Aggregated seed arrival alters plant diversity in grassland communities

Aims Species aggregation is commonly seen in plant communities and may increase diversity by causing intraspecific competition to exceed interspecific competition. One potential source of this spatial aggregation is seed dispersal but it is unclear to what extent aggregated seed distributions affect plant diversity in real communities. Using a field experiment, I tested whether uniform or aggregated seed arrival alters community structure and whether these effects vary with sowing density.Methods The experiment consisted of two spatial seeding treatments (uniform and aggregated) that were fully crossed with three seed density treatments. Sixty, 3 × 4-m plots were arrayed in a low-diversity grassland located in Kansas, USA. Each plot was divided into forty-eight, 0.5 × 0.5-m patches. For aggregated seeding treatments, each of the 15 species was sown into three randomly selected patches within the plot (3×15 = 45). To create a uniform species arrival but control for the seed addition method, all 15 species were sown into 45 individual patches (with three patches remaining unsown) within each plot. Seed mass for each species was held constant at the plot scale between uniform or aggregated treatments within a given level of the sowing density treatment. After two growing seasons, plant density was quantified for all sown species in 15 randomly selected patches from each plot.Important findings I found evidence for shifts in community structure in response to the different spatial seeding patterns. The evenness of added species was higher under aggregated than uniform sowing patterns. There was no detectable effect of aggregated seed sowing on species richness at 3.75 m 2 scale. However, when species richness was extrapolated to larger scales (11.25 m 2), aggregated sowing was predicted to have greater richness than uniform sowing. Effects of seed aggregation on community structure were apparent only at moderate to high sowing rates, yet the latter are within the range of measured seed dispersal in similar grasslands. Additionally, as sowing density increased, seed mass became an increasingly effective predictor of relative abundances for added species, but only under uniform sowing patterns supporting the idea that aggregated dispersal may buffer weaker (smaller seeded) species from competition during colonization. This is the first experiment to show that aggregated seed dispersal patterns can increase at least some components of plant diversity in undisturbed grasslands and suggests that previous seed dispersal experiments, which utilize uniform seed sowing, may underestimate the potential effect of dispersal on plant community structure.     

Plant density can increase invertebrate postdispersal seed predation in an experimental grassland community

Janzen–Connell effects are negative effects on the survival of a plant's progeny at high conspecific densities or close to its conspecifics. Although the role of Janzen–Connell effects on the maintenance of plant diversity was frequently studied, only few studies targeted Janzen–Connell effects via postdispersal seed predation in temperate grassland systems. We examined effects of conspecific density (abundance of conspecific adult plants) on postdispersal seed predation by invertebrates of three grassland species (Centaurea jacea, Geranium pratense, and Knautia arvensis) in experimental plant communities. Additionally, we examined the impact of plant species richness and different seed predator communities on total and relative seed predation (= seed predation of one plant species relative to others). We offered seeds in an exclusion experiment, where treatments allowed access for (1) arthropods and slugs, (2) arthropods only, (3) small arthropods only, and (4) slugs only. Treatments were placed in plots covering a gradient of abundance of conspecific adults at different levels of plant species richness (1, 2, 3, 4, 8 species). Two of the plant species (C. jacea and K. arvensis) experienced higher rates of seed predation and relative predation with increasing abundance of conspecific adults. For C. jacea, this effect was mitigated with increasing plant species richness. Differences in seed predator communities shifted seed predation between the plant species and changed the magnitude of seed predation of one plant species relative to the others. We exemplify density‐dependent increase in seed predation via invertebrates in grassland communities shaping both the total magnitude of species‐specific seed predation and seed predation of one species relative to others. Further differences in seed predator groups shift the magnitude of seed predation between different plant species. This highlights the importance of invertebrate seed predation to structure grasslands via density‐dependent effects and differing preferences of consumer groups.     

Control of plant species diversity and community invasibility by species immigration: seed richness versus seed density

Immigration rates of species into communities are widely understood to influence community diversity, which in turn is widely expected to influence the susceptibility of ecosystems to species invasion. For a given community, however, immigration processes may impact diversity by means of two separable components: the number of species represented in seed inputs and the density of seed per species. The independent effects of these components on plant species diversity and consequent rates of invasion are poorly understood. We constructed experimental plant communities through repeated seed additions to independently measure the effects of seed richness and seed density on the trajectory of species diversity during the development of annual plant communities. Because we sowed species not found in the immediate study area, we were able to assess the invasibility of the resulting communities by recording the rate of establishment of species from adjacent vegetation. Early in community development when species only weakly interacted, seed richness had a strong effect on community diversity whereas seed density had little effect. After the plants became established, the effect of seed richness on measured diversity strongly depended on seed density, and disappeared at the highest level of seed density. The ability of surrounding vegetation to invade the experimental communities was decreased by seed density but not by seed richness, primarily because the individual effects of a few sown species could explain the observed invasion rates. These results suggest that seed density is just as important as seed richness in the control of species diversity, and perhaps a more important determinant of community invasibility than seed richness in dynamic plant assemblages.     

鼎湖山植物群落多样性的研究

根据鼎湖山自然保护区１６个植物群落的样地调查资料,从种丰富度和不同类型的物种多样性指数、均匀度与植被类型、人为干扰程度、演替阶段等方面的关系进行了分析。本区植被类型的多样性指数、均匀率指数变化反映了其结构复杂程度、生境的差异。物种丰富和多样性随着人为干扰程度的增强而降低。群落内各层的物种丰富度的大小的顺序为“灌木层＞草本层＞乔木层”,而群落多样性指数的大小顺序为“灌木层＞乔木层＞草本层”。群落多样     

Relationships between arthropod richness,evenness, and diversity are altered by complementarity among plant genotypes

Biodiversity is quantified via richness (e.g., the number of species), evenness (the relative abundance distribution of those species), or proportional diversity (a combination of richness and evenness, such as the Shannon index, H′). While empirical studies show no consistent relationship between these aspects of biodiversity within communities, the mechanisms leading to inconsistent relationships have received little attention. Here, using common evening primrose (Oenothera biennis) and its associated arthropod community, we show that relationships between arthropod richness, evenness, and proportional diversity are altered by plant genotypic richness. Arthropod richness increased with O. biennis genotypic richness due to an abundance-driven accumulation of species in response to greater plant biomass. Arthropod evenness and proportional diversity decreased with plant genotypic richness due to a nonadditive increase in abundance of a dominant arthropod, the generalist florivore/omnivore Plagiognathas politus (Miridae). The greater quantity of flowers and buds produced in polycultures—which resulted from positive complementarity among O. biennis genotypes—increased the abundance of this dominant insect. Using choice bioassays, we show that floral quality did not change in plant genotypic mixtures. These results elucidate mechanisms for how plant genotypic richness can modify relationships between arthropod richness, evenness, and proportional diversity. More broadly, our results suggest that trophic interactions may be a previously underappreciated factor controlling relationships between these different aspects of biodiversity.     

Sowing density effects and patterns of colonization in a prairie restoration

A cost‐effective approach in plant restorations could be to increase sowing density for species known to be challenging to establish, while reducing sowing density for species that easily colonize on their own. Sowing need not occur evenly across the site for rapidly dispersing species. We explored these issues using a prairie restoration experiment on a high‐school campus with three treatments: plots sown only to grasses (G plots), to grasses and forbs (GF1), and to grasses and forbs with forbs sown at twice the density (GF2). In year 2, GF1 and GF2 plots had higher diversity than G plots, as expected, but GF2 treatments did not have twice the sown forb cover. However, high forb sowing density increased forb richness, probably by reducing stochastic factors in establishment. Cover of nonsown species was highest in G plots and lowest in GF2 plots, suggesting suppressive effects of native forbs on weedy species. Colonization of G plots by two sown forbs (Coreopsis tinctoria and Rudbeckia hirta) was apparent after 2.5 years, providing evidence that these species are self‐sustaining. Colonization was greater in edges than in the central areas of G plots. Through construction of establishment kernels, we infer that the mean establishment distance was shorter for R. hirta (6.7 m) compared to C. tinctoria (21.1 m). Our results lead us to advocate for restoration practices that consider not only seed sowing but also subsequent dispersal of sown species. Furthermore, we conclude that restoration research is particularly amenable for outdoor education and university‐high school collaborations.     

庙岛群岛典型植物群落物种、功能、结构多样性及其对环境因子的响应

海岛植被在全球生物多样性研究中起重要作用,研究海岛植被多样性对于理解海陆相互作用下植物群落的多样性维持机制有重要意义.本研究以庙岛群岛的麻栎群落、刺槐群落、黑松群落、荆条群落4种典型植物群落为对象,采用物种多样性指数、功能多样性指数和结构多样性指数,在群落尺度上探讨了海岛典型植物群落物种、功能、结构多样性间的关系及其对环境因子的响应.结果表明: 黑松群落的物种丰富度与Rao指数高于刺槐群落与麻栎群落,而结构多样性却较低;荆条灌丛的物种、结构多样性均低于森林群落,而功能多样性高于部分森林群落.物种丰富度与Rao指数以及树高多样性间呈显著正相关,与功能均匀度呈显著负相关.结构多样性主要由坡度决定且与坡度呈负相关;功能均匀度与坡度呈正相关,而功能异质性、功能离散度和物种多样性则更多地受土壤理化性质的影响,与土壤容重及土壤总碳呈正相关,与土壤含水率呈负相关.总体而言,庙岛群岛的植物群落多样性格局既有与大陆植被相似的特征,但也有其海岛特殊性.     

Warming increases plant biomass and reduces diversity across continents,latitudes, and species migration scenarios in experimental wetland communities

Atmospheric warming may influence plant productivity and diversity and induce poleward migration of species, altering communities across latitudes. Complicating the picture is that communities from different continents deviate in evolutionary histories, which may modify responses to warming and migration. We used experimental wetland plant communities grown from seed banks as model systems to determine whether effects of warming on biomass production and species richness are consistent across continents, latitudes, and migration scenarios. We collected soil samples from each of three tidal freshwater marshes in estuaries at three latitudes (north, middle, south) on the Atlantic coasts of Europe and North America. In one experiment, we exposed soil seed bank communities from each latitude and continent to ambient and elevated (+2.8 °C) temperatures in the greenhouse. In a second experiment, soil samples were mixed either within each estuary (limited migration) or among estuaries from different latitudes in each continent (complete migration). Seed bank communities of these migration scenarios were also exposed to ambient and elevated temperatures and contrasted with a no‐migration treatment. In the first experiment, warming overall increased biomass (+16%) and decreased species richness (?14%) across latitudes in Europe and North America. Species richness and evenness of south‐latitude communities were less affected by warming than those of middle and north latitudes. In the second experiment, warming also stimulated biomass and lowered species richness. In addition, complete migration led to increased species richness (+60% in North America, + 100% in Europe), but this higher diversity did not translate into increased biomass. Species responded idiosyncratically to warming, but Lythrum salicaria and Bidens sp. increased significantly in response to warming in both continents. These results reveal for the first time consistent impacts of warming on biomass and species richness for temperate wetland plant communities across continents, latitudes, and migration scenarios.     

Patterns of Plant Species Diversity in Remnant and Restored Tallgrass Prairies

To restore diversity of native vegetation, we must understand factors responsible for diversity in targeted communities. These factors operate at different spatial scales and may affect the number and relative abundances of species differently. We measured diversity of plant species and functional groups of species in replicated plots within paired restored and remnant (relic) tallgrass prairies at three locations in central Texas, U.S.A. To determine the contributions of species abundances and of spatial patterns of diversity to differences between prairie types, we separated diversity into richness and evenness (relative biomass) and into within‐plot (α), among‐plot (β), and prairie (γ) components. Species diversity was greater in remnant than in restored prairies at all spatial scales. At the γ scale, both species richness and species evenness were greater in remnants because of greater spatial variation in species composition. At the α scale, remnants were more diverse because of greater richness alone. Mean α richness correlated positively with the size of the species pool in restored prairies only, implying that in remnants, α richness was influenced more by colonization dynamics than by the number of species available for colonization. Plots in remnant prairies contained more functional groups and fewer species per group than did plots in restored prairies, suggesting that resource partitioning was greater in relic prairies. Our results are consistent with the interpretation that local ecological processes, like resource partitioning and limitations on seed dispersal, contribute to the greater diversity of remnant than restored prairies in central Texas. Restoration practices that limit abundances of competitive dominants, increase the number of species in seed mixtures, and increase the proximity of plants of different functional groups thus may be required to better simulate the plant diversity of tallgrass prairies.     

Negative relationships between species richness and evenness render common diversity indices inadequate for assessing long-term trends in butterfly diversity

Species richness and evenness, the two principle components of species diversity, are frequently used to describe variation in species assemblages in space and time. Compound indices, including variations of both the Shannon–Wiener index and Simpson’s index, are assumed to intelligibly integrate species richness and evenness into all-encompassing measures. However, the efficacy of compound indices is disputed by the possibility of inverse relationships between species richness and evenness. Past studies have assessed relationships between various diversity measures across survey locations for a variety of taxa, often finding species richness and evenness to be inversely related. Butterflies are one of the most intensively monitored taxa worldwide, but have been largely neglected in such studies. Long-term butterfly monitoring programs provide a unique opportunity for analyzing how trends in species diversity relate to habitat and environmental conditions. However, analyzing trends in butterfly diversity first requires an assessment of the applicability of common diversity measures to butterfly assemblages. To accomplish this, we quantified relationships between butterfly diversity measures estimated from 10 years of butterfly population data collected in the North Saskatchewan River Valley in Edmonton, Alberta, Canada. Species richness and evenness were inversely related within the butterfly assemblage. We conclude that species evenness may be used in conjunction with richness to deepen our understandings of assemblage organization, but combining these two components within compound indices does not produce measures that consistently align with our intuitive sense of species diversity.     

Should I plant or should I sow? Restoration outcomes compared across seven riparian revegetation projects

To revegetate native plant communities, it is often cheaper to direct seed than to plant nursery‐grown stock. However, the outcomes of direct seeding can be quite variable, and it is unclear whether direct seeding or planting is more likely to facilitate the restoration of diverse plant communities. To address this question, we compared the outcomes of each method across several recent riparian revegetation projects where both direct seeding and tube‐stock planting were used. We surveyed riparian revegetation projects at seven sites within the greater Melbourne area that had been revegetated between 1 and 4 years previously. Sites were all on land previously used for agriculture or degraded public land and ranged in environmental and climatic conditions. Woody plant density, establishment of target species, species richness, species diversity (evenness) and plant heights were assessed. Direct seeding tended to result in higher plant densities and similar species richness, but lower rates of species establishment and diversity compared with planting. A median of 67% of target species established via direct seeding compared with 100% for planting, with direct seeded areas often dominated by one or two species. In general, overall revegetation outcomes were often driven by climatic and site factors, rather than revegetation method. We suggest that to achieve good restoration outcomes from revegetation in riparian areas, a bet‐hedging or combined approach using both sowing and planting may be the best strategy.     

山西翅果油树群落的多样性研究

用丰富度指数、多样性指数和均匀度指数对山西翅果油树群落的多样性进行了研究 ,并用相关分析研究了海拔与多样性指数及多样性指数间的关系 ,结果表明 :1 )在干扰强烈的生境中 ,翅果油树群落具有较低的丰富度指数、多样性指数的均匀性 ,而接近顶极群落阶段 ,多样性指数和丰富度指数也较低 ,但具有较高的均匀性 ;干扰强度较小的生境中 ,群落具有较高的丰富度指数、多样性指数和均匀性。2 )灌木层和草本层的丰富度、多样性指数和均匀度指数呈现多元化的趋势。 3)海拔对山西翅果油树群落多样性的影响不显著。     

Richness, structure and functioning in metazoan parasite communities

Ecosystem functioning, characterized by components such as productivity and stability, has been extensively linked with diversity in recent years, mainly in plant ecology. The aim of our study was thus to quantify general relationships between diversity, community structure and ecosystem functions in metazoan parasite communities. We used data on parasite communities from 15 species of marine fish hosts from coastal Chile. The volumetric abundance (volume of all parasite species per individual host, in mm³) was used as a surrogate for productivity. Species diversity was measured using both species richness and evenness, while community structure was estimated using the co‐occurrence indices V‐ratio, C‐score and a new C‐score_s index standardized for the number of host replicates. After correcting for fish size, 47% of host species show no relationship, 13% show a hump shaped curve and 40% show positive monotonic relationships between productivity and parasite richness across all host individuals in a sample. We obtained a logarithmically decreasing relationship between evenness and productivity for all fish species, and propose a ‘dominance‐resistance’ hypothesis based on immunity to explain this pattern. The stability of the parasite community, measured as the coefficient of variation in productivity among individual hosts, was strongly and positively related to mean species richness across the 15 host species. The C‐score_s index, based on the number of checkerboard units in the host‐parasite presence/absence matrix, increases linearly with mean productivity across the 15 host species, suggesting that parasite communities tend to be more structured when they are more productive. This is the likely reason why linear relationships between richness and productivity were not observed consistently in all fish species. Parasite communities provide some clear patterns for the diversity–ecosystem functioning debate in ecology, although other factors, such as the history of community assembly, may also influence these patterns.     

山西五鹿山森林群落木本植物功能多样性

通过选取群落中木本植物种子的扩散方式、传粉方式、植株高度和盖度等13个功能性状,计算出群落的6个功能多样性指数:功能性状距离、功能性状平均距离、功能体积、功能均匀度、功能分散指数和Rao二次熵指数,结合群落物种丰富度指数、Shannon-Wiener指数和物种均匀度指数对山西五鹿山森林群落木本植物功能多样性进行研究。结果表明:(1)功能性状距离、功能性状平均距离、功能体积与物种丰富度、Shannon-Wiener指数显著正相关;功能均匀度与Shannon-Wiener指数、物种均匀度指数显著正相关;功能分散指数、Rao二次熵指数与物种均匀度指数、Shannon-Wiener指数显著正相关;(2)功能多样性的差异很大程度上是由于物种差异所引起的;(3)6个功能多样性指数可分为三类:功能性状距离、功能性状平均距离、功能体积为功能丰富度指数;功能均匀度为功能均匀度指数;功能分散指数和Rao二次熵指数为功能离散度指数。该分类结果符合指数的计算方法和生态学意义,以及相互独立的标准。     

The Abundance of Large Ateline Monkeys is Positively Associated with the Diversity of Plants Regenerating in Neotropical Forests

Ateline monkeys, the largest primates in the Neotropics, may disperse more than one million seeds/km²/d at sites where they are abundant, but it is unclear whether a reduction in their populations can alter plant diversity patterns. The species richness and composition of regenerating plants as a proxy of future plant communities were studied by comparing 16 sites with different ateline abundance in three countries in northwestern South America. A total of 3658 plots included 94,340 regenerating plants, which were assigned to species or morphospecies. Paired t‐tests comparing sites in the same region but with different densities of atelines, and regression analyses showed a consistent positive relationship between ateline density and plant diversity. These results were due to the larger number of stems per area and higher evenness at sites with more atelines, suggesting higher recruitment rates for dispersed seeds. Differences were also found in plant composition, as canopy, endozoochorous, and medium seed size plants were consistently more abundant in sites with more ateline monkeys than in sites with less atelines. The findings of this study suggest that these primates play a key role in plant regeneration. In order to maintain the diversity and plant composition of tropical forests for future generations, conservation of these large frugivores and other key game species is imperative.     

佛坪国家级自然保护区植物群落物种多样性特征

用物种多样性指数（Simpson指数和Shannon指数）、均匀度指数（Pielou均匀度）和物种丰富度,对佛坪国家级自然保护区主要高等植物群落物种多样性进行了测度。结果表明：森林群落的多样性程度高于灌丛和草甸,落叶阔叶林高于针叶林,栎林高于杨桦林。群落物种多样性有随海拔高度的升高而降低的趋势。物种多样性在森林群落垂直结构上的分布特点,表现为一般乔木层与灌木层接近但低于草本层。     

Consistent Effects of Biodiversity on Ecosystem Functioning Under Varying Density and Evenness

Biodiversity experiments typically vary only species richness and composition, yet the generality of their results relies on consistent effects of these factors even under varying starting conditions of density and evenness. We tested this assumption in a factorial species richness x density x evenness experiment using a pool of 60 common grassland species divided into four functional groups (grasses, legumes, tall herbs and short herbs). Richness varied from 1, 2, 4, 8 to 16 species, total planting density was 1,000 or 2,000 seeds/m², and species were sown in even or uneven proportions, where one functional group was made dominant. Aboveground plant biomass increased linearly with the logarithm of species richness in all density and evenness treatments during all three years of the experiment. This was due to a convergence of realized density and evenness within species richness levels, although functional groups which were initially made dominant retained their dominance. Between species richness levels, realized density increased, and realized evenness decreased with species richness. Thus, more individuals could coexist if they belonged to different species. Within species richness levels, higher biomass values were correlated with lower density, suggesting an underlying thinning process. However, communities with low realized evenness also had low biomass values; thus high biomass did not result from species dominance. So-called complementarity and selection effects were similar across density and evenness treatments, indicating that the mechanisms underpinning the biodiversity effects were not altered. Species richness was the dominant driver of aboveground biomass, irrespective of variations in total densities and species abundance distributions at the start of the experiment; rejecting the hypothesis that initial differences in species abundance distributions might lead to different “stable states” in community structure or biomass. Thus, results from previous biodiversity experiments that only manipulated species richness and composition should be quite robust and broadly generalizable.     

When competition does not matter: grassland diversity and community composition

We examined whether the intense root competition in a rough fescue grassland plant community in central Alberta, Canada, was important in structuring plant species diversity or community composition. We measured competition intensity across gradients of species richness, evenness, and community composition, using pairs of naturally occurring plants of 12 species. One plant in each pair was isolated from neighbors to measure competition; community structure and environmental conditions were also measured at each pair. We used structural equation modeling to examine how competition influenced community structure. Competition intensity was unrelated to species richness and community composition, but increased competition intensity was associated with a slight decline in evenness. Size-symmetric root competition was probably unimportant in structuring this plant community because there are no feedback mechanisms through which size-symmetric competition can magnify small initial differences and eventually lead to competitive exclusion. In plant communities with little shoot competition, competition and community structure should be unlinked regardless of competition intensity. In more productive systems, we propose that interactions between root and shoot competition may indirectly structure communities by altering the overall asymmetry of competition.     

Reductions in grassland species evenness increase dicot seedling invasion and spittle bug infestation

Previous experiments that tested whether diverse plant communities have lower invasibility have all varied species richness. We experimentally varied evenness of four grassland species (three grasses and one forb) by planting a field experiment in Texas, and monitored the number of unplanted dicot and monocot species that invaded plots for two growing seasons. By varying evenness, we eliminated any sampling effect in our diversity treatment, because all plots contained the same plant species. Experimentally reducing evenness led to a greater number of dicot invaders, which emerged in plots throughout the growing season, but had less of an effect on monocot invaders, which emerged in flushes when experimental plants were semi‐dormant. Frequency of Solidago canadensis (altissima) stems with spittle bugs significantly increased with reductions in evenness during the first year, apparently because the greater number of Solidago stems in high evenness plots diluted the spittle‐bug effect. These results support the view that higher diversity plant communities are more resistant to dicot invaders and insect herbivores.