Nutrient enrichment increased species richness of leaf litter fungal assemblages in a tropical forest

Microbial communities play a major role in terrestrial ecosystem functioning, but the determinates of their diversity and functional interactions are not well known. In this study, we explored leaf litter fungal diversity in a diverse Panama lowland tropical forest in which a replicated factorial N, P, K and micronutrient fertilization experiment of 40 × 40 m plots had been ongoing for nine years. We extracted DNA from leaf litter samples and used fungal‐specific amplification and a 454 pyrosequencing approach to sequence two loci, the nuclear ribosomal internal transcribed spacer (ITS) region and the nuclear ribosomal large subunit (LSU) D1 region. Using a 95% sequence similarity threshold for ITS1 spacer recovered a total of 2523 OTUs, and the number of unique ITS1 OTUs per 0.5–1.0 g leaf litter sample ranged from 55 to 177. Ascomycota were the dominant phylum among the leaf litter fungi (71% of the OTUs), followed by Basidiomycota (26% of the OTUs). In contrast to our expectations based on temperate ecosystems, long‐term addition of nutrients increased, rather than decreased, species richness relative to controls. Effect of individual nutrients was more subtle and seen primarily as changes in community compositions especially at lower taxonomic levels, rather than as significant changes in species richness. For example, plots receiving P tended to show a greater similarity in community composition compared to the other nutrient treatments, the +PK, +NK and +NPK plots appeared to be more dominated by the Nectriaceae than other treatments, and indicator species for particular nutrient combinations were identified.     

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity–stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity–stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non‐native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi‐natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.     

Increasing synchrony opposes stabilizing effects of species richness on terrestrial communities

Aim

Ecological theory has predicted that species richness should stabilize communities, with mechanisms including species synchrony and population variability determining the net impacts. While these theories have been supported empirically, results can be sensitive to taxonomic bias as studies are often focussed on plants. Trophic differences between consumers and primary producers can lead to varying stabilizing effects of species richness. Here, we compared the impact of species richness on community variability in four taxonomic groups: terrestrial birds, mammals, invertebrates and plants.

Location

Global.

Method

Using data from 6763 time series globally (BioTIME) for four terrestrial taxa, we quantified community and population variability and species synchrony based on abundance fluctuations over time.

Results

Species richness destabilized communities through increasing synchrony and stabilized communities through reducing population variability in all taxa. Such opposing effects weakened the net impacts of species richness on communities. Population variability had higher importance relative to synchrony in plant communities. By contrast, synchrony had more comparable (or even higher) importance compared with population variability in animal communities. When synchrony and population variability were not controlled, stabilizing impacts of species richness were detected in plant communities only.

Main Conclusions

Our results highlight how species richness drives stabilizing and destabilizing mechanisms simultaneously across all taxa, with strong taxonomic variation in the relative importance of these mechanisms in regulating community variability. This questions the generality of previous findings on stabilizing impacts of species richness based on limited taxonomic coverage. Additionally, our results indicate the need to understand how the importance of stabilizing and destabilizing mechanisms differs in determining community variability across organisms and environments.     

Grazing weakens temporal stabilizing effects of diversity in the Eurasian steppe

Many biodiversity experiments have demonstrated that plant diversity can stabilize productivity in experimental grasslands. However, less is known about how diversity–stability relationships are mediated by grazing. Grazing is known for causing species losses, but its effects on plant functional groups (PFGs) composition and species asynchrony, which are closely correlated with ecosystem stability, remain unclear. We conducted a six‐year grazing experiment in a semi‐arid steppe, using seven levels of grazing intensity (0, 1.5, 3.0, 4.5, 6.0, 7.5, and 9.0 sheep per hectare) and two grazing systems (i.e., a traditional, continuous grazing system during the growing period (TGS), and a mixed one rotating grazing and mowing annually (MGS)), to examine the effects of grazing system and grazing intensity on the abundance and composition of PFGs and diversity–stability relationships. Ecosystem stability was similar between mixed and continuous grazing treatments. However, within the two grazing systems, stability was maintained through different pathways, that is, along with grazing intensity, persistence biomass variations in MGS, and compensatory interactions of PFGs in their biomass variations in TGS. Ecosystem temporal stability was not decreased by species loss but rather remain unchanged by the strong compensatory effects between PFGs, or a higher grazing‐induced decrease in species asynchrony at higher diversity, and a higher grazing‐induced increase in the temporal variation of productivity in diverse communities. Ecosystem stability of aboveground net primary production was not related to species richness in both grazing systems. High grazing intensity weakened the temporal stabilizing effects of diversity in this semi‐arid grassland. Our results demonstrate that the productivity of dominant PFGs is more important than species richness for maximizing stability in this system. This study distinguishes grazing intensity and grazing system from diversity effects on the temporal stability, highlighting the need to better understand how grazing regulates ecosystem stability, plant diversity, and their synergic relationships.     

Nitrogen enrichment weakens ecosystem stability through decreased species asynchrony and population stability in a temperate grassland

Biodiversity generally promotes ecosystem stability. To assess whether the diversity–stability relationship observed under ambient nitrogen (N) conditions still holds under N enriched conditions, we designed a 6‐year field experiment to test whether the magnitude and frequency of N enrichment affects ecosystem stability and its relationship with species diversity in a temperate grassland. Results of this experiment showed that the frequency of N addition had no effect on either the temporal stability of ecosystem and population or the relationship between diversity and stability. Nitrogen addition decreased ecosystem stability significantly through decreases in species asynchrony and population stability. Species richness was positively associated with ecosystem stability, but no significant relationship between diversity and the residuals of ecosystem stability was detected after controlling for the effects of the magnitude of N addition, suggesting collinearity between the effects of N addition and species richness on ecosystem stability, with the former prevailing over the latter. Both population stability and the residuals of population stability after controlling for the effects of the magnitude of N addition were positively associated with ecosystem stability, indicating that the stabilizing effects of component populations were still present after N enrichment. Our study supports the theory predicting that the effects of environmental factors on ecosystem functioning are stronger than those of biodiversity. Understanding such mechanisms is important and urgent to protect biodiversity in mediating ecosystem functioning and services in the face of global changes.     

An empirical test of the mass effect determinant of species richness

The mass effect determinant of species richness proposed by Shmida & Wilson (1985) was empirically tested at the interface of undisturbed, native vegetation with recovering vegetation on a reclaimed surface mine. The observed decrease in native species richness with distance away from the source area was consistent with the predicted pattern based on Shmida & Wilson's (1985) model.Taxonomic nomenclature follows Hitchcock & Cronquist (1983) except genus Artemisia, which follows Beetle & Johnson (1982).This research was supported by Grant No. DEB81-01827 from the National Science Foundation. Neil West and Charles Romesburg aided in the experimental design and analysis. Patricia Johnson assisted in computer programming.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.     

Nutrient stoichiometry and land use rather than species richness determine plant functional diversity

Plant functional traits reflect individual and community ecological strategies. They allow the detection of directional changes in community dynamics and ecosystemic processes, being an additional tool to assess biodiversity than species richness. Analysis of functional patterns in plant communities provides mechanistic insight into biodiversity alterations due to anthropogenic activity. Although studies have consi‐dered of either anthropogenic management or nutrient availability on functional traits in temperate grasslands, studies combining effects of both drivers are scarce. Here, we assessed the impacts of management intensity (fertilization, mowing, grazing), nutrient stoichiometry (C, N, P, K), and vegetation composition on community‐weighted means (CWMs) and functional diversity (Rao's Q) from seven plant traits in 150 grasslands in three regions in Germany, using data of 6 years. Land use and nutrient stoichiometry accounted for larger proportions of model variance of CWM and Rao's Q than species richness and productivity. Grazing affected all analyzed trait groups; fertilization and mowing only impacted generative traits. Grazing was clearly associated with nutrient retention strategies, that is, investing in durable structures and production of fewer, less variable seed. Phenological variability was increased. Fertilization and mowing decreased seed number/mass variability, indicating competition‐related effects. Impacts of nutrient stoichiometry on trait syndromes varied. Nutrient limitation (large N:P, C:N ratios) promoted species with conservative strategies, that is, investment in durable plant structures rather than fast growth, fewer seed, and delayed flowering onset. In contrast to seed mass, leaf‐economics variability was reduced under P shortage. Species diversity was positively associated with the variability of generative traits. Synthesis. Here, land use, nutrient availability, species richness, and plant functional strategies have been shown to interact complexly, driving community composition, and vegetation responses to management intensity. We suggest that deeper understanding of underlying mechanisms shaping community assembly and biodiversity will require analyzing all these parameters.     

The mid-domain effect: geometric constraints on the geography of species richness

Geographic patterns of species richness are influenced by many factors, but the role of shared physiographical and physiological boundaries in relation to range-size distributions has been surprisingly neglected, in spite of the fact that such geometric constraints lead to mid-domain richness peaks even without environmental gradients (the mid-domain effect). Relying on null models, several recent studies have begun to quantify this problem using simulated and empirical data. This approach promises to transform how we perceive geographic variation in diversity, including the long unresolved latitudinal gradient in species richness. The question is not whether geometry affects such patterns, but by how much.     

Experimental climate change weakens the insurance effect of biodiversity

Ecosystems are simultaneously affected by biodiversity loss and climate change, but we know little about how these factors interact. We predicted that climate warming and CO (2) -enrichment should strengthen trophic cascades by reducing the relative efficiency of predation-resistant herbivores, if herbivore consumption rate trades off with predation resistance. This weakens the insurance effect of herbivore diversity. We tested this prediction using experimental ocean warming and acidification in seagrass mesocosms. Meta-analyses of published experiments first indicated that consumption rate trades off with predation resistance. The experiment then showed that three common herbivores together controlled macroalgae and facilitated seagrass dominance, regardless of climate change. When the predation-vulnerable herbivore was excluded in normal conditions, the two resistant herbivores maintained top-down control. Under warming, however, increased algal growth outstripped control by herbivores and the system became algal-dominated. Consequently, climate change can reduce the relative efficiency of resistant herbivores and weaken the insurance effect of biodiversity.     

Effects of woody plant species richness on mammal species richness in southern Africa

Aim To determine the relationship between the species richness of woody plants and that of mammals after accounting for the effect of environmental variables. Location Southern Africa, including Namibia, South Africa, Lesotho, Swaziland, Botswana, Zimbabwe, and part of Mozambique. Methods We used a comprehensive dataset including the species richness of mammals and of woody plants and environmental variables for 118 quadrats (each of 25,000 km²) across southern Africa, and used structural equation models (SEMs) and spatial regressions to examine the relationship between the species richness of woody plants and of mammal trophic guilds (herbivores, insectivores, carni/omnivores) and habitat guilds (aquatic/fossorial, ground‐living, climbers, aerial), after controlling for environment. We compared the results of SEMs with those of single‐predictor regressions (without controlling for environment) and of spatial regressions (controlling for both environment and residual spatial autocorrelation). Results The geographical variation of mammal species richness in southern Africa was strongly and positively related to that of woody plant species richness, and this relationship held for most mammal guilds even when the influence of environment and spatial autocorrelation had been accounted for. However, the effect of woody plant species richness on the richness of aquatic/fossorial species almost disappeared after controlling for environment, suggesting that the congruence in species richness patterns between these two groups results from similar responses to the same environmental variables. For many mammal guilds, the relative role of environmental predictors as measured by standardized partial regression coefficients changed depending on whether non‐spatial single‐predictor regressions, non‐spatial SEMs, or spatial regressions were used. Main conclusions Woody plants are important determinants of the species richness of most mammal guilds in southern Africa, even when controlling for environment and residual spatial autocorrelation. Environmental correlates with animal species richness as measured by simple correlations or single‐predictor regressions might not always reflect direct effects; they might, at least to some degree, result from indirect effects via woody plants. Interpretations of the strength of the effect of environmental variables on mammal species richness in southern Africa depend largely on whether spatial or non‐spatial models are used. We therefore stress the need for caution when interpreting environmental ‘effects’ on broad‐scale patterns of species richness if spatial and non‐spatial methods yield contrasting results.     

Plant species richness under

Exotic pine plantations constitute a significant landscape feature in the North Island of New Zealand but their conservation value for native plant species is not often documented. Pine stem density, height and basal area of nine plantations of Pinus radiata ranging in age from 6 to 67 years in Kinleith Forest was determined. Pines reached heights of 60 m, and stand basal areas up to 183 ± 14 m(2)ha(-1). The abundance of woody shrubs, tree ferns and ground ferns was assessed in each stand. Understorey composition of shrubs and ferns was reflected on the first two axes of DCA ordinations and correlated with the age of the pines. Adventive shrubs predominated in stands < 20 years old. Light-demanding native shrubs with bird dispersed fruits predominated in older stands, with more shade-tolerant species in the oldest site. Species richness increased rapidly in the first 11 years, but thereafter more slowly. Twelve native shrub species and 22 ferns were recorded from the most diverse stands. Richness and species composition were related to stand age, and probably also to topographical heterogeneity and aspect. Tree ferns reached densities of 2000—2500 ha(-1) and basal areas of 20—30 m(2)ha(-1) in the older stands. Initially the tree fern population was strongly dominated by Dicksonia squarrosa, which comprised 84% of individuals overall. Five species were present by 29 years. The faster growing Cynthea medullaris and C. smithii achieved greater heights than the Dicksonia spp., and their relative biomass was greatest in the oldest stands.     

Ciliate community associated with aquatic macrophyte roots: effects of nutrient enrichment on the community composition and species richness

The objective of this study was to identify the impact of nutrient enrichment on the diversity of the ciliate community associated with the roots of the aquatic macrophyte Eichhornia crassipes. The experiment was performed in the Garças Lake, located in the Upper Paraná River floodplain, Brazil. We conducted two treatments (fertilized and control) with three replicates each. To increase the initial nutrient concentrations in each mesocosm of the fertilized treatment, we added 1000 μg L⁻¹ of KNO₃ and 200 μg L⁻¹ of KH₂PO₄ during each sampling date. We found a relative high number of ciliate species (85 species) and a predominance of hypotrichs. Among the recorded species, about 25% occurred exclusively in the fertilized treatment. Moreover, detrended correspondence analysis demonstrated that the ciliate community associated with E. crassipes roots changed significantly in response to the nutrient input in such a way that the species composition of the fertilized treatment was remarkably different from that of the control. In contrast to our expectations, species richness in the fertilized treatment was significantly higher than that in the control, refuting our hypothesis that species richness decreases under eutrophic conditions.     

The effect of naturally varying discharge rates on the species richness of lotic midges

Species richness significantly changed both temporally, over a two year period, and spatially, across three sites in a sandy run, in an assemblage of aquatic insects (Chironomidae) inhabiting a fourth order stream in southeastern Ohio, USA. A total of 25 species was encountered, and for a given site and time, richness varied between 0 and 13 species/site with a mean of 3.3. Averaged over two years, the species diversity trend from bank to channel was an increasing number of species with a decreasing variance, indicating a greater stability in species richness in the channel than near the bank. A significant amount of the temporal variability in richness can be explained by variation in the stream discharge rate. Discharge was inversely related to a) species richness and b) the rate of change of species richness, for two sites in the stream center but not for a more protected site near the bank. Abugov's (1982) model of the phasing of disturbances may explain the species richness patterns observed in this study.     

Neutral community dynamics, the mid-domain effect and spatial patterns in species richness

The mid‐domain effect (MDE) aims to explain spatial patterns in species richness invoking only stochasticity and geometrical constraints. In this paper, we used simulations to show that its main qualitative prediction, a hump‐shaped pattern in species richness, converges to the expectation of a spatially bounded neutral model when communities are linked by short‐distance migration. As these two models can be linked under specific situations, neutral theory may provide a mechanistic population level basis for MDE. This link also allows establishing in which situations MDE patterns are more likely to be found. Also, in this situation, MDE models could be used as a first approximation to understand the role of both stochastic (ecological drift and migration) and deterministic (adaptation to environmental conditions) processes driving the spatial structure of species richness.     

Edge effect of a pine plantation reduces dry grassland invertebrate species richness

Natural steppes in European agricultural landscapes are characterized by high biotic richness but are subject to fragmentation and associated edge effects. Edge effects on species richness were investigated at an ecotone from a pine plantation to a short-grass steppe in Eastern Austria for eleven invertebrate taxa differentiated into habitat guilds based on known live-history strategies of individual species (grassland species, forest species, generalist species), including Red-Listed and non-threatened grassland species. The large size of the studied grassland site provided an opportunity to test edge effects in the absence of confounding factors and to a gradient length of 208 m into the grassland habitat along a clear-cut border to a pine plantation. All sampling was done by pitfall trapping. Species richness of habitat guilds, but not total richness, was effectively explained by biotic variables reflecting the influence of shading in particular (i.e. soil temperature sums). Total species richness showed a bimodal response pattern, with increases towards the habitat edge and interior grassland habitat. Habitat guilds showed diverging responses to distances from the edge, but no saturation in species richness, with a continuum of edge effects across the entire distance of the grassland samples. Our findings contrast those of previous investigations based on samples taken from smaller patches and across shorter distances from the edge. Methodological and conservation implications are discussed.     

Independence of total species richness and richness difference are not the same thing

Leprieur and Oikonomou (2014) criticized that a replacement index β_− 3, a partitioned component of Jaccard index β_jac, was not richness independent and should not be used in biogeographic and ecological studies. However, Leprieur and Oikonomou failed to recognize the difference between richness and richness difference. Independence of total species richness is not equal to independence of richness difference. Theoretically and ideally, it is true that β_− 3 (and β_jac as well) is not independent of richness difference while Simpson index (β_sim) is fully independent of richness difference. However, all these indices actually are independent of total species richness. At last, it is worth mentioning that the ideal independence studied here is easily violated in computational simulation and real-world settings.     

Nutrient enrichment can increase the severity of coral diseases

The prevalence and severity of marine diseases have increased over the last 20 years, significantly impacting a variety of foundation and keystone species. One explanation is that changes in the environment caused by human activities have impaired host resistance and/or have increased pathogen virulence. Here, we report evidence from field experiments that nutrient enrichment can significantly increase the severity of two important Caribbean coral epizootics: aspergillosis of the common gorgonian sea fan Gorgonia ventalina and yellow band disease of the reef‐building corals Montastraea annularis and M. franksii. Experimentally increasing nutrient concentrations by 2–5× nearly doubled host tissue loss caused by yellow band disease. In a separate experiment, nutrient enrichment significantly increased two measures of sea fan aspergillosis severity. Our results may help explain the conspicuous patchiness of coral disease severity, besides suggesting that minimizing nutrient pollution could be an important management tool for controlling coral epizootics.