Scale-dependent responses of plant biodiversity to nitrogen enrichment

Experimental studies demonstrating that nitrogen (N) enrichment reduces plant diversity within individual plots have led to the conclusion that anthropogenic N enrichment is a threat to global biodiversity. These conclusions overlook the influence of spatial scale, however, as N enrichment may alter beta diversity (i.e., how similar plots are in their species composition), which would likely alter the degree to which N-induced changes in diversity within localities translate to changes in diversity at larger scales that are relevant to policy and management. Currently, it is unclear how N enrichment affects biodiversity at scales larger than a small plot. We synthesized data from 18 N-enrichment experiments across North America to examine the effects of N enrichment on plant species diversity at three spatial scales: small (within plots), intermediate (among plots), and large (within and among plots). We found that N enrichment reduced plant diversity within plots by an average of 25% (ranging from a reduction of 61% to an increase of 5%) and frequently enhanced beta diversity. The extent to which N enrichment altered beta diversity, however, varied substantially among sites (from a 22% increase to an 18% reduction) and was contingent on site productivity. Specifically, N enrichment enhanced beta diversity at low-productivity sites but reduced beta diversity at high-productivity sites. N-induced changes in beta diversity generally reduced the extent of species loss at larger scales to an average of 22% (ranging from a reduction of 54% to an increase of 18%). Our results demonstrate that N enrichment often reduces biodiversity at both local and regional scales, but that a focus on the effects of N enrichment on biodiversity at small spatial scales may often overestimate (and sometimes underestimate) declines in regional biodiversity by failing to recognize the effects of N on beta diversity.     

Increasing enemy biodiversity strengthens herbivore suppression on two plant species

Concern over biodiversity loss, especially at higher trophic levels, has led to a surge in studies investigating how changes in natural enemy diversity affect community and ecosystem functioning. These studies have found that increasing enemy diversity can strengthen, weaken, and not affect prey suppression, demonstrating that multi-enemy effects on prey are context-dependent. Here we ask how one factor, plant species identity, influences multi-enemy effects on prey. We focused on two plant species of agricultural importance, potato (Solanum tuberosum), and collards (Brassica oleracea L.). These species share a common herbivorous pest, the green peach aphid (Myzus persicae), but vary in structural and chemical traits that affect aphid reproductive rates and which may also influence inter-enemy interactions. In a large-scale field experiment, overall prey exploitation varied dramatically among the plant species, with enemies reducing aphid populations by approximately 94% on potatoes and approximately 62% on collards. Increasing enemy diversity similarly strengthened aphid suppression on both plants, however, and there was no evidence that plant species identity significantly altered the relationship between enemy diversity and prey suppression. Microcosm experiments suggested that, on both collards and potatoes, intraspecific competition among natural enemies exceeded interspecific competition. Enemy species showed consistent and significant differences in where they foraged on the plants, and enemies in the low-diversity treatment tended to spend less time foraging than enemies in the high-diversity treatment. These data suggest that increasing enemy diversity may strengthen aphid suppression because interspecific differences in where enemies forage on the plant allow for greater resource partitioning. Further, these functional benefits of diversity appear to be robust to changes in plant species identity.     

Hierarchical spatial point process analysis for a plant community with high biodiversity

A complex multivariate spatial point pattern of a plant community with high biodiversity is modelled using a hierarchical multivariate point process model. In the model, interactions between plants with different post-fire regeneration strategies are of key interest. We consider initially a maximum likelihood approach to inference where problems arise due to unknown interaction radii for the plants. We next demonstrate that a Bayesian approach provides a flexible framework for incorporating prior information concerning the interaction radii. From an ecological perspective, we are able both to confirm existing knowledge on species’ interactions and to generate new biological questions and hypotheses on species’ interactions.

外来入侵杂草空心莲子草对植物生物多样性的影响

利用样方法调查了南京市空心莲子草(Alternantheraphiloxeroides)生长生境中杂草的种类和分布。结果表明,南京市秋季空心莲子草分布样点有杂草30科101种,其中以禾本科、菊科、蓼科居多。主成分分析表明,影响杂草分布的主要因素是土壤水分条件和人为干扰强度。经Q型分析, 24个样点可划分为3个聚类群:聚类群Ⅰ包括的样点均分布在路边和田边开阔地以及小河、小沟和湖的岸边,以马唐、狗尾草和牛筋草为优势种;聚类群Ⅱ的样点都是农田隔离带如田埂以及农田中的沟壑,以波斯婆婆纳、碎米莎草、野塘蒿和铁苋菜为优势种;聚类群Ⅲ的样点以菜地和果园为主,以马唐、香附子、马齿苋和黄鹌菜为优势种。分析了空心莲子草重要值与物种丰富度之间的关系,当空心莲子草重要值大于1. 5时,样方物种丰富度随重要值的增加而减小。2者之间具有显著相关性,表明空心莲子草的入侵对生物多样性有不利影响。     

Relationship between plant biodiversity and heavy metal bioavailability in grasslands overlying an abandoned mine

Abandoned metal mines in the Sierra de Guadarrama, Madrid, Spain, are often located in areas of high ecological value. This is true of an abandoned barium mine situated in the heart of a bird sanctuary. Today the area sustains grasslands, interspersed with oakwood formations of Quercus ilex and heywood scrub (Retama sphaerocarpa L.), used by cattle, sheep and wild animals. Our study was designed to establish a relationship between the plant biodiversity of these grasslands and the bioavailability of heavy metals in the topsoil layer of this abandoned mine. We conducted soil chemical analyses and performed a greenhouse evaluation of the effects of different soil heavy metal concentrations on biodiversity. The greenhouse bioassays were run for 6 months using soil samples obtained from the mine polluted with heavy metals (Cu, Zn, Pb and Cd) and from a control pasture. Soil heavy metal and Na concentrations, along with the pH, had intense negative effects on plant biodiversity, as determined through changes in the Shannon index and species richness. Numbers of grasses, legumes, and composites were reduced, whilst other species (including ruderals) were affected to a lesser extent. Zinc had the greatest effect on biodiversity, followed by Cd and Cu. When we compared the sensitivity of the biodiversity indicators to the different metal content variables, pseudototal metal concentrations determined by X-ray fluorescence (XRF) were the most sensitive, followed by available and soluble metal contents. Worse correlations between biodiversity variables and metal variables were shown by pseudototal contents obtained by plasma emission spectroscopy (ICP-OES). Our results highlight the importance of using as many different indicators as possible to reliably assess the response shown by plants to heavy metal soil pollution.     

Ecological drivers of group living in two populations of the communally rearing rodent, Octodon degus

Intraspecific variation in sociality is thought to reflect a trade-off between current fitness benefits and costs that emerge from individuals' decision to join or leave groups. Since those benefits and costs may be influenced by ecological conditions, ecological variation remains a major, ultimate cause of intraspecific variation in sociality. Intraspecific comparisons of mammalian sociality across populations facing different environmental conditions have not provided a consistent relationship between ecological variation and group-living. Thus, we studied two populations of the communally rearing rodent Octodon degus to determine how co-variation between sociality and ecology supports alternative ecological causes of group living. In particular, we examined how variables linked to predation risk, thermal conditions, burrowing costs, and food availability predicted temporal and population variation in sociality. Our study revealed population and temporal variation in total group size and group composition that covaried with population and yearly differences in ecology. In particular, predation risk and burrowing costs are supported as drivers of this social variation in degus. Thermal differences, food quantity and quality were not significant predictors of social group size. In contrast to between populations, social variation within populations was largely uncoupled from ecological differences.     

Longevity can buffer plant and animal populations against changing climatic variability

Both means and year-to-year variances of climate variables such as temperature and precipitation are predicted to change. However, the potential impact of changing climatic variability on the fate of populations has been largely unexamined. We analyzed multiyear demographic data for 36 plant and animal species with a broad range of life histories and types of environment to ask how sensitive their long-term stochastic population growth rates are likely to be to changes in the means and standard deviations of vital rates (survival, reproduction, growth) in response to changing climate. We quantified responsiveness using elasticities of the long-term population growth rate predicted by stochastic projection matrix models. Short-lived species (insects and annual plants and algae) are predicted to be more strongly (and negatively) affected by increasing vital rate variability relative to longer-lived species (perennial plants, birds, ungulates). Taxonomic affiliation has little power to explain sensitivity to increasing variability once longevity has been taken into account. Our results highlight the potential vulnerability of short-lived species to an increasingly variable climate, but also suggest that problems associated with short-lived undesirable species (agricultural pests, disease vectors, invasive weedy plants) may be exacerbated in regions where climate variability decreases.     

Simulation of small bird populations. II. Reduced breeding in two British raptor species

A computer model was developed to simulate the fate of small populations of birds. It uses general and easily available data as input. Monte-Carlo techniques are used and a survival probability is calculated for every population member four times per simulated year. The model allows for density-dependence in winter survival and also in fecundity.Simulation results are used to compile standard age-specific life-tables for all complete cohorts generated. A mean life-table is also made. The survivor functions, both accumulated and mean, are contrasted with control functions and tested for significance.The model has been applied to classical population data in ornithology, namely those for the great tit and the tawny owl in Wytham Woods (Oxford, UK). There was a fair agreement between simulation results and field data.     

Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

Increased plant size in exotic populations: a common-garden test with 14 invasive species

The "evolution of increased competitive ability" (EICA) hypothesis predicts that exotic species will adapt to reduced herbivore pressure by losing costly defenses in favor of competitive ability. Previous studies often support the prediction that plants from exotic populations will be less well defended than plants from native populations. However, results are mixed with respect to the question of whether plants from exotic populations have become more competitive. In a common-garden experiment involving plants from two native and two exotic populations of 14 different invasive species, we tested whether exotic plants generally grow larger than conspecific native plants, and whether patterns of relative growth depend on the intensity of competition. We found a quite consistent pattern of larger exotic than native plants, but only in the absence of competition. These results suggest that invasive species may often evolve increased growth, and that increased growth may facilitate adaptation to noncompetitive environments.     

论生物多样性的保护问题

当前，任何地区的生物多样性都是自然和社会因素长期综合作用所造成。生物多样性最丰富的地区不一定全部由顶极植被所覆盖，而常常是一些包括所有演替类型的地方。本文通过讨论自然环境的变化和人类生产活动对不同区域景观的影响，探讨有关生物多样性的保护问题。     

Predictability of marine nematode biodiversity

In this paper, we investigated: (1) the predictability of different aspects of biodiversity, (2) the effect of spatial autocorrelation on the predictability and (3) the environmental variables affecting the biodiversity of free-living marine nematodes on the Belgian Continental Shelf. An extensive historical database of free-living marine nematodes was employed to model different aspects of biodiversity: species richness, evenness, and taxonomic diversity. Artificial neural networks (ANNs), often considered as “black boxes”, were applied as a modeling tool. Three methods were used to reveal these “black boxes” and to identify the contributions of each environmental variable to the diversity indices. Since spatial autocorrelation is known to introduce bias in spatial analyses, Moran's I was used to test the spatial dependency of the diversity indices and the residuals of the model. The best predictions were made for evenness. Although species richness was quite accurately predicted as well, the residuals indicated a lack of performance of the model. Pure taxonomic diversity shows high spatial variability and is difficult to model. The biodiversity indices show a strong spatial dependency, opposed to the residuals of the models, indicating that the environmental variables explain the spatial variability of the diversity indices adequately. The most important environmental variables structuring evenness are clay and sand fraction, and the minimum annual total suspended matter. Species richness is also affected by the intensity of sand extraction and the amount of gravel of the sea bed.     

A nearly neutral model of biodiversity

S. P. Hubbell's unified neutral theory of biodiversity has stimulated much new thinking about biodiversity. However, empirical support for the neutral theory is limited, and several observations are inconsistent with the predictions of the theory, including positive correlations between traits associated with competitive ability and species abundance and correlations between species diversity and ecosystem functioning. The neutral theory can be extended to explain these observations by allowing species to differ slightly in their competitive ability (fitness). Here, we show that even slight differences in fecundity can greatly reduce the time to extinction of competitors even when the community size is large and dispersal is spatially limited. In this case, species richness is dramatically reduced, and a markedly different species abundance distribution is predicted than under pure neutrality. In the nearly neutral model, species co-occur in the same community not because of, but in spite of, ecological differences. The more competitive species with higher fecundity tend to have higher abundance both in the metacommunity and in local communities. The nearly neutral perspective provides a theoretical framework that unites the sampling model of the neutral theory with theory of biodiversity affecting ecosystem function.     

Large-scale biodiversity patterns in freshwater phytoplankton

Our planet shows striking gradients in the species richness of plants and animals, from high biodiversity in the tropics to low biodiversity in polar and high-mountain regions. Recently, similar patterns have been described for some groups of microorganisms, but the large-scale biogeographical distribution of freshwater phytoplankton diversity is still largely unknown. We examined the species diversity of freshwater phytoplankton sampled from 540 lakes and reservoirs distributed across the continental United States and found strong latitudinal, longitudinal, and altitudinal gradients in phytoplankton biodiversity, demonstrating that microorganisms can show substantial geographic variation in biodiversity. Detailed analysis using structural equation models indicated that these large-scale biodiversity gradients in freshwater phytoplankton diversity were mainly driven by local environmental factors, although there were residual direct effects of latitude, longitude, and altitude as well. Specifically, we found that phytoplankton species richness was an increasing saturating function of lake chlorophyll a concentration, increased with lake surface area and possibly increased with water temperature, resembling effects of productivity, habitat area, and temperature on diversity patterns commonly observed for macroorganisms. In turn, these local environmental factors varied along latitudinal, longitudinal, and altitudinal gradients. These results imply that changes in land use or climate that affect these local environmental factors are likely to have major impacts on large-scale biodiversity patterns of freshwater phytoplankton.     

Not just the usual suspects: insect herbivore populations and communities are associated with multiple plant nutrients

The relationship between plant nutrient content and insect herbivore populations and community structure has long interested ecologists. Insect herbivores require multiple nutrients, but ecologists have focused mostly on nitrogen (an estimate of plant protein content), and more recently phosphorus (P); other nutrients have received little attention. Here we document nutrient variation in grass and forb samples from grassland habitats in central Nebraska using an elemental approach; in total we measured foliar concentrations of 12 elements (N and P, plus S, B, Ca, Mg, Na, K, Zn, Fe, Mn, and Cu). We detected significant variability among sites for N, P, Mg, Na, K, and Cu. We next used a model selection approach to explore how this nutritional variation and plant biomass correlate with grasshopper densities (collectively and at the feeding-guild level), and principal component analysis to explore nutrient correlations with grasshopper community species composition. When all grasshoppers were pooled, densities varied among sites, but only P was associated with abundance of the elements shown to vary between sites. Different responses occurred at the feeding-guild level. For grass specialists, densities were associated with N, plus P, Mg, and Na. For forb specialists, N and P were often associated with density, but associations with Na and K were also observed. Finally, mixed-feeder abundance was strongly associated with biomass, and to a lesser extent P, Mg, Na, and Cu. At the community level, B, Ca, Zn, and Cu, plus biomass, explained > 30% of species composition variation. Our results confirm the positive association of N and P with insect herbivore populations, while suggesting a potential role for Mg, Na, and K. They also demonstrate the importance of exploring effects at the feeding-guild level. We hope our data motivate ecologists to think beyond N and P when considering plant nutrient effects on insect herbivores, and make a call for studies to examine functional responses of insect herbivores to dietary manipulation of Mg, Na, and K. Finally, our results demonstrate correlations between variation in nutrients and species assemblages, but factors not linked to plant nutrient quality or biomass likely explain most of the observed variation.     

Modeling biodiversity dynamics in countryside landscapes

The future of biodiversity hinges to a great extent on the conservation value of countryside, the growing fraction of Earth's surface heavily influenced by human activities. How many species, and which species, can persist in such landscapes (and analogous seascapes) are open questions. Here we explore two complementary theoretical frameworks to address these questions: species-area relationships and demographic models. We use the terrestrial mammal fauna of Central America to illustrate the application of both frameworks. We begin by proposing a multi-habitat species-area relationship, the countryside species-area relationship, to forecast species extinction rates. To apply it, we classify the mammal fauna by affinity to native and human-dominated habitats. We show how considering the conservation value of countryside habitats changes estimates derived from the classic species-area approach We also examine how the z value of the species-area relationship affects extinction estimates. Next, we present a framework for assessing the relative vulnerability of species to extinction in the countryside, based on the Skellam model of population dynamics. This model predicts the minimum area of contiguous native habitat required for persistence of a species, which we use as an indicator of vulnerability to habitat change. To apply the model, we use our habitat affinity classification of mammals and we estimate life-history parameters by species and habitat type. The resulting ranking of vulnerabilities is significantly correlated with the World Conservation Union (IUCN) Red List assessment.     

Foliar fungal pathogens and grassland biodiversity

By attacking plants, herbivorous mammals, insects, and belowground pathogens are known to play an important role in maintaining biodiversity in grasslands. Foliar fungal pathogens are ubiquitous in grassland ecosystems, but little is known about their role as drivers of community composition and diversity. Here we excluded foliar fungal pathogens from perennial grassland by using fungicide to determine the effect of natural levels of disease on an otherwise undisturbed plant community. Importantly, we excluded foliar fungal pathogens along with rabbits, insects, and mollusks in a full factorial design, which allowed a comparison of pathogen effects along with those of better studied plant enemies. This revealed that fungal pathogens substantially reduced aboveground plant biomass and promoted plant diversity and that this especially benefited legumes. The scale of pathogen effects on productivity and biodiversity was similar to that of rabbits and insects, but different plant species responded to the exclusion of the three plant enemies. These results suggest that theories of plant coexistence and management of biodiversity in grasslands should consider foliar fungal pathogens as potentially important drivers of community composition.     

Effect of streambed substrate on macroinvertebrate biodiversity

Macroinvertebrates are important components of stream ecosystems, and are often used as indicator species for the assessment of river ecology. Numerous studies have shown that substrate is the primary physical environmental variable affecting the taxa richness and density of macroinvertebrates. The aim of this work is to study the effects of the characteristics of streambed substrate, such as grain size, shape, and roughness, on the composition and biodiversity of macroinvertebrates. A field experiment was done on the Juma River, a second-order mountain stream in northern China. Substrata of cobbles, hewn stones, pebbles, coarse sand, and fine sand were used to replace the original gravel and sand bed in a stretch of 30 m in length. The sampling results indicated that the macroinvertebrate assemblage is significantly affected by the grain size, porosity and interstitial dimension of the substrate, while it is rarely affected by the shape and the surface roughness of the experimental substrata. Macroinvertebrate compositions in cobbles and hewn stones were stable and changed least over time. The taxa richness and density of individuals in the substrata of cobbles, hewn stones, and pebbles are much higher than in those of the coarse sand and fine sand.     

Global biodiversity patterns of benthic marine algae

Species richness patterns are remarkably similar across many marine taxa, yet explanations of how such patterns are generated and maintained are conflicting. I use published occurrence data to identify previously masked latitudinal and longitudinal diversity gradients for all genera of benthic marine macroalgae and for species in the Order Bryopsidales. I also quantify the size, location, and overlap of macroalgal geographic ranges to determine how the observed richness patterns are generated. Algal genera exhibit an inverse latitudinal gradient, with biodiversity hotspots in temperate regions, while bryopsidalean species reach peak diversity in the tropics. The geographic distribution of range locations results in distinct clusters of range mid-points. In particular, widespread taxa are centered within tight latitudinal and longitudinal bands in the middle of the Indo-Pacific and Atlantic Oceans while small-ranged taxa are clustered in peripheral locations, suggesting that variation in speciation and extinction are important drivers of algal diversity patterns. Hypotheses about factors that regulate diversity contain underlying assumptions about the size and location of geographic ranges, in addition to predictions as to why species numbers will differ among regions. Yet these assumptions are rarely considered in assessing the validity of the prevailing hypotheses. I assess a suite of hypotheses, suggested to explain patterns of marine diversity, by comparing algal-richness patterns in combination with the size and location of algal geographic ranges, to the richness and range locations predicted by these hypotheses. In particular, the results implicate habitat areas and ocean currents as the most plausible drivers of observed diversity patterns.