Saturating effects of species diversity on life-history evolution in bacteria

Species interactions can play a major role in shaping evolution in new environments. In theory, species interactions can either stimulate evolution by promoting coevolution or inhibit evolution by constraining ecological opportunity. The relative strength of these effects should vary as species richness increases, and yet there has been little evidence for evolution of component species in communities. We evolved bacterial microcosms containing between 1 and 12 species in three different environments. Growth rates and yields of isolates that evolved in communities were lower than those that evolved in monocultures, consistent with recent theory that competition constrains species to specialize on narrower sets of resources. This effect saturated or reversed at higher levels of richness, consistent with theory that directional effects of species interactions should weaken in more diverse communities. Species varied considerably, however, in their responses to both environment and richness levels. Mechanistic models and experiments are now needed to understand and predict joint evolutionary dynamics of species in diverse communities.     

Plant history and soil history jointly influence the selection environment for plant species in a long‐term grassland biodiversity experiment

Long‐term biodiversity experiments have shown increasing strengths of biodiversity effects on plant productivity over time. However, little is known about rapid evolutionary processes in response to plant community diversity, which could contribute to explaining the strengthening positive relationship. To address this issue, we performed a transplant experiment with offspring of seeds collected from four grass species in a 14‐year‐old biodiversity experiment (Jena Experiment). We used two‐ and six‐species communities and removed the vegetation of the study plots to exclude plant–plant interactions. In a reciprocal design, we transplanted five “home” phytometers (same origin and actual environment), five “away‐same” phytometers (same species richness of origin and actual environment, but different plant composition), and five “away‐different” phytometers (different species richness of origin and actual environment) of the same species in the study plots. In the establishment year, plants transplanted in home soil produced more shoots than plants in away soil indicating that plant populations at low and high diversity developed differently over time depending on their associated soil community and/or conditions. In the second year, offspring of individuals selected at high diversity generally had a higher performance (biomass production and fitness) than offspring of individuals selected at low diversity, regardless of the transplant environment. This suggests that plants at low and high diversity showed rapid evolutionary responses measurable in their phenotype. Our findings provide first empirical evidence that loss of productivity at low diversity is not only caused by changes in abiotic and biotic conditions but also that plants respond to this by a change in their micro‐evolution. Thus, we conclude that eco‐evolutionary feedbacks of plants at low and high diversity are critical to fully understand why the positive influence of diversity on plant productivity is strengthening through time.     

Soil microbiome mediates positive plant diversity‐productivity relationships in late successional grassland species

Which processes drive the productivity benefits of biodiversity remain a critical, but unanswered question in ecology. We tested whether the soil microbiome mediates the diversity‐productivity relationships among late successional plant species. We found that productivity increased with plant richness in diverse soil communities, but not with low‐diversity mixtures of arbuscular mycorrhizal fungi or in pasteurised soils. Diversity‐interaction modelling revealed that pairwise interactions among species best explained the positive diversity‐productivity relationships, and that transgressive overyielding resulting from positive complementarity was only observed with the late successional soil microbiome, which was both the most diverse and exhibited the strongest community differentiation among plant species. We found evidence that both dilution/suppression from host‐specific pathogens and microbiome‐mediated resource partitioning contributed to positive diversity‐productivity relationships and overyielding. Our results suggest that re‐establishment of a diverse, late successional soil microbiome may be critical to the restoration of the functional benefits of plant diversity following anthropogenic disturbance.     

Architectural design influences the diversity and structure of the built environment microbiome

Buildings are complex ecosystems that house trillions of microorganisms interacting with each other, with humans and with their environment. Understanding the ecological and evolutionary processes that determine the diversity and composition of the built environment microbiome—the community of microorganisms that live indoors—is important for understanding the relationship between building design, biodiversity and human health. In this study, we used high-throughput sequencing of the bacterial 16S rRNA gene to quantify relationships between building attributes and airborne bacterial communities at a health-care facility. We quantified airborne bacterial community structure and environmental conditions in patient rooms exposed to mechanical or window ventilation and in outdoor air. The phylogenetic diversity of airborne bacterial communities was lower indoors than outdoors, and mechanically ventilated rooms contained less diverse microbial communities than did window-ventilated rooms. Bacterial communities in indoor environments contained many taxa that are absent or rare outdoors, including taxa closely related to potential human pathogens. Building attributes, specifically the source of ventilation air, airflow rates, relative humidity and temperature, were correlated with the diversity and composition of indoor bacterial communities. The relative abundance of bacteria closely related to human pathogens was higher indoors than outdoors, and higher in rooms with lower airflow rates and lower relative humidity. The observed relationship between building design and airborne bacterial diversity suggests that we can manage indoor environments, altering through building design and operation the community of microbial species that potentially colonize the human microbiome during our time indoors.     

Species interactions alter evolutionary responses to a novel environment

Studies of evolutionary responses to novel environments typically consider single species or perhaps pairs of interacting species. However, all organisms co-occur with many other species, resulting in evolutionary dynamics that might not match those predicted using single species approaches. Recent theories predict that species interactions in diverse systems can influence how component species evolve in response to environmental change. In turn, evolution might have consequences for ecosystem functioning. We used experimental communities of five bacterial species to show that species interactions have a major impact on adaptation to a novel environment in the laboratory. Species in communities diverged in their use of resources compared with the same species in monocultures and evolved to use waste products generated by other species. This generally led to a trade-off between adaptation to the abiotic and biotic components of the environment, such that species evolving in communities had lower growth rates when assayed in the absence of other species. Based on growth assays and on nuclear magnetic resonance (NMR) spectroscopy of resource use, all species evolved more in communities than they did in monocultures. The evolutionary changes had significant repercussions for the functioning of these experimental ecosystems: communities reassembled from isolates that had evolved in polyculture were more productive than those reassembled from isolates that had evolved in monoculture. Our results show that the way in which species adapt to new environments depends critically on the biotic environment of co-occurring species. Moreover, predicting how functioning of complex ecosystems will respond to an environmental change requires knowing how species interactions will evolve.     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Species richness, environmental fluctuations, and temporal change in total community biomass

Theory and empirical results suggest that high biodiversity should often cause lower temporal variability in aggregate community properties such as total community biomass. We assembled microbial communities containing 2 to 8 species of competitors in aquatic microcosms and found that the temporal change in total community biomass was positively but insignificantly associated with diversity in a constant temperature environment. There was no evidence of any trend in variable temperature environments. Three non-exclusive mechanisms might explain the lack of a net stabilising effect of species richness on temporal change. (1) A direct destabilising effect of diversity on population level variances caused some populations to vary more when embedded in more diverse communities. (2) Similar responses of the different species to environmental variability might have limited any insurance effect of increased species richness. (3) Large differences in the population level variability of different species (i.e., unevenness) could weaken the relation between species richness and community level stability. These three mechanisms may outweigh the stabilising effects of increases in total community biomass with diversity, statistical averaging, and slightly more negative covariance in more diverse communities. Our experiment and analyses advocate for further experimental investigations of diversity-variability relations.     

Long-term stability in the richness and structure of helminth communities in eels,Anguilla anguilla,in Lough Derg,River Shannon,Ireland

A data set on intestinal helminth parasites was collected in the course of an 18 year investigation into the biology of eels in Meelick Bay, Lough Derg, River Shannon. This was used to test two hypotheses relating to the composition and structure of intestinal helminth communities, namely that eels in large rivers do not harbour richer and more diverse communities than those in small rivers but that community composition and structure are more stable over time than in small rivers. The helminth community was species poor, with only six species comprising the component community and a maximum infracommunity richness of three species. The community was overwhelmingly dominated by the acanthocephalan Acanthocephalus lucii, reflecting the importance of its intermediate host Asellus aquaticus in the eels' diet. The remaining helminth species contributed to species richness but made very little contribution to community diversity. Population levels of Acanthocephalus lucii fell and remained low between 1992 and 2000, probably reflecting increased movement of eels from other parts of the lough into Meelick Bay. Diversity values were low, but similar to those reported from other rivers in Britain and Europe. The results provided support for both hypotheses and indicated that in respect of richness, diversity and dominance, the helminth communities of eels in the River Shannon were typical of, and comparable to, those of other large rivers throughout Europe.     

Soil Communities Promote Temporal Stability and Species Asynchrony in Experimental Grassland Communities

Background

Over the past two decades many studies have demonstrated that plant species diversity promotes primary productivity and stability in grassland ecosystems. Additionally, soil community characteristics have also been shown to influence the productivity and composition of plant communities, yet little is known about whether soil communities also play a role in stabilizing the productivity of an ecosystem.

Methodology/Principal Findings

Here we use microcosms to assess the effects of the presence of soil communities on plant community dynamics and stability over a one-year time span. Microcosms were filled with sterilized soil and inoculated with either unaltered field soil or field soil sterilized to eliminate the naturally occurring soil biota. Eliminating the naturally occurring soil biota not only resulted in lower plant productivity, and reduced plant species diversity, and evenness, but also destabilized the net aboveground productivity of the plant communities over time, which was largely driven by changes in abundance of the dominant grass Lolium perenne. In contrast, the grass and legumes contributed more to net aboveground productivity of the plant communities in microcosms where soil biota had been inoculated. Additionally, the forbs exhibited compensatory dynamics with grasses and legumes, thus lowering temporal variation in productivity in microcosms that received the unaltered soil inocula. Overall, asynchrony among plant species was higher in microcosms where an unaltered soil community had been inoculated, which lead to higher temporal stability in community productivity.

Conclusions/Significance

Our results suggest that soil communities increase plant species asynchrony and stabilize plant community productivity by equalizing the performance among competing plant species through potential antagonistic and facilitative effects on individual plant species.     

Resilience of the natural arthropod community on apple to external disturbance

Abstract.

• 1 A naturally evolved arthropod community in a 6-year-old apple orchard was treated with three applications of permethrin; a second naturally evolved community was studied as an untreated control. Disturbance to the community was measured with Shannon's index of species diversity for the phytophagous and beneficial portions of the community.
• 2 Initially, there was a reduction in diversity of both phytophagous and beneficial arthropods because of the insecticide. Reduction in diversity was a result of both lower number of species and lower evenness of species abundance.
• 3 Two months after the last permethrin spray, there was no difference between diversity in the phytophagous community, but the beneficial community was more diverse in the treated orchard than in the untreated control.
• 4 The year after treatment there were few differences between the phytophagous communities, but the beneficial community was more diverse in the treated orchard than in the untreated orchard in May and early June; however, by September the beneficial community was less diverse in the treated orchard.
• 5 Although diversity statistics of the phytophagous communities were similar 15 months after treatment, differences still remained with the treated community being dominated by more r-selected species and the control orchard dominated by more K-selected species.
• 6 The arthropod community on apple has a high level of stability as reflected by its resilience to a severe external disturbance. This stability would allow for large, but infrequent, disturbances for pest management and still maintain long-term ecological equilibrium in the community.

     

Mycorrhizal fungal spore community structure in a manipulated prairie

Most plant communities support a diverse assemblage of arbuscular mycorrhizal fungi (AMF). AMF communities have the potential to affect plant community structure and vice versa. We examined AMF sporulation in a 4.5‐ha reconstructed prairie in Eau Claire County, Wisconsin. In fall 2003, the site was planted with varied numbers and combinations of native prairie species from four functional guilds: C₃ grasses/sedges, C₄ grasses, legume, and nonleguminous forbs. We hypothesized that more diverse plant seeding mixtures would promote AMF diversity. To examine the interaction between plant and fungal communities, plots were divided and subplots treated with the fungicide chlorothalonil to suppress AMF, enriched with ammonium nitrate fertilizer, treated with both fungicide and nitrogen, or remained untreated (control). Soil samples were collected during the summers of 2004, 2006, and 2007 from each subplot. Spores of AMF were extracted, identified to species, and enumerated. Initial plant seeding diversity did not significantly influence spore abundance, fungal diversity, plant productivity, or plant richness 4 years after establishment. Fungal species richness was positively, but weakly, correlated with plant productivity (r² = 0.11) and plant richness (r² = 0.09). Fungal community composition changed significantly over time; nitrogen addition, fungicide application, and site characteristics also shaped community composition. After 4 years of treatment, nitrogen and fungicide reduced AMF richness, changed sporulation patterns among AMF taxa, and reduced diversity and productivity in plant communities. Divergence in AMF community is being mirrored by changes in the plant community independent of initial seeding treatments, though causation could not be determined.     

A unifying evolutionary theory for the biomass–diversity–fertility relationship

Although a widely accepted ecological theory predicts that more diverse plant communities should be better able to capture resources and turn carbon dioxide into biomass, the most productive communities known are low diversity agricultural ones. This paradox has fuelled a long running controversy in ecology surrounding the nature of the relationship between diversity, productivity and fertility. Here, an evolutionary computer model is used which demonstrates that given the opportunity, species-rich communities may evolve under high fertility conditions. In contrast to low diversity, highly productive agricultural communities are shown to probably be a recent phenomenon. In simulations where fertility was applied to communities that had evolved under lower nutrient conditions, a few species had the ability to become ‘dominant’. These species were responsible for the loss of diversity and for the majority of biomass production. These results are consistent with complementarity theory applying in nature in old co-evolved low nutrient communities, whereas in recently established fertile agricultural communities, dominant species appear to regulate biomass production. Understanding the nature of these ‘dominant’ species throws light on our understanding of phenotypic plasticity and the ecology of invasive species. The appendices files are currently available at .     

Genetic diversity affects productivity in early but not late stages of stand development

Recent declines in the genetic diversity of populations have stimulated research on the importance of genetic diversity for the functioning of natural communities. Current studies on this topic are based on the exploration of a limited number of clones and do not allow distinctions to be made between the effects of genetic identity and genetic diversity per se and to evaluate the effects of genetic diversity in genetically diverse communities. Also, most information comes from short-term studies, which are insufficient for evaluating the long-term effects relevant in relatively undisturbed communities of perennial species.We explored the importance of clone diversity vs. clone identity for stand productivity and the changes of the pattern over time. We used 18 clones of a perennial grass, Festuca rubra, to establish a set of communities composed of 1, 6 or 18 clones in two environments and studied the effects of genetic diversity on stand productivity over 3 years.Genetic diversity had a significant effect on stand productivity in the 1st year but not in the 2nd or 3rd year. In most cases, the observed yield was not significantly different from the total expected yield. The biomass of the mixtures never outperformed the biomass of the most productive clone, suggesting that clone identity is an important determinant of total biomass.The results indicate that the effects of genetic diversity on stand productivity may be transient and suggest that the conclusions of short-term studies on diversity effects should be evaluated carefully. They also suggest that individual clones are not complementary and that the properties of the stands are mainly additive results of the properties of the constituent clones.     

Soil fertility increases with plant species diversity in a long-term biodiversity experiment

Most explanations for the positive effect of plant species diversity on productivity have focused on the efficiency of resource use, implicitly assuming that resource supply is constant. To test this assumption, we grew seedlings of Echinacea purpurea in soil collected beneath 10-year-old, experimental plant communities containing one, two, four, eight, or 16 native grassland species. The results of this greenhouse bioassay challenge the assumption of constant resource supply; we found that bioassay seedlings grown in soil collected from experimental communities containing 16 plant species produced 70% more biomass than seedlings grown in soil collected beneath monocultures. This increase was likely attributable to greater soil N availability, which had increased in higher diversity communities over the 10-year-duration of the experiment. In a distinction akin to the selection/complementarity partition commonly made in studies of diversity and productivity, we further determined whether the additive effects of functional groups or the interactive effects of functional groups explained the increase in fertility with diversity. The increase in bioassay seedling biomass with diversity was largely explained by a concomitant increase in N-fixer, C4 grass, forb, and C3 grass biomass with diversity, suggesting that the additive effects of these four functional groups at higher diversity contributed to enhance N availability and retention. Nevertheless, diversity still explained a significant amount of the residual variation in bioassay seedling biomass after functional group biomass was included in a multiple regression, suggesting that interactions also increased fertility in diverse communities. Our results suggest a mechanism, the fertility effect, by which increased plant species diversity may increase community productivity over time by increasing the supply of nutrients via both greater inputs and greater retention.     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

草甸和沼泽植物群落功能多样性与生产力的关系

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Effects of different components of diversity on productivity in artificial plant communities

In an experiment on artificial plant communities, the effects of three components of plant diversity—plant species diversity, plant functional group diversity and plant functional diversity—on community productivity and soil water content were compared. We found that simple regression analysis showed a positive diversity effect on ecosystem processes (productivity and soil water content). However, when three components of diversity were included in the multiple regression analyses, the results showed that functional group diversity and functional diversity had more important effects on productivity and resource use efficiency. These results suggested that, compared with species number, functional differences among species and the range of functional traits carried by plants are the basis of biodiversity effects on ecosystem functioning. These diversity effects of increasing functional group diversity or functional diversity were likely because species differing greatly in size, life form, phenology and capacity to capture and use resources efficiently in diverse communities realize complementary resource use in temporal, spatial, and biological ways.     

Dynamics, diversity, and resource gradient relationships in the herbaceous layer of an old-growth Appalachian forest

The ecological drivers of herbaceous layer composition and diversity in deciduous forests of eastern North America are imperfectly understood. We analyzed the herbaceous layer, across the growing season, in a central Appalachian old-growth forest to examine dynamics, diversity, and relationships to resource gradients. We found clear variation in herb species composition over the growing season. We identified intermingled resource gradients, including soil nutrients, light availability, and topography, that were related to herbaceous composition. We found that herb layer diversity was different among previously identified tree communities, but was not variable over the growing season. We identified a unimodal relationship between diversity and productivity in the herb flora that held throughout the growing season despite changing composition and levels of productivity. Diversity and distributions in the herbaceous community of our study site are linked to a complex of resource gradients.     

Relationship between functional diversity and productivity in meadow and marsh plant communities

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Mutualistic rhizobia reduce plant diversity and alter community composition

Mutualistic interactions can be just as important to community dynamics as antagonistic species interactions like competition and predation. Because of their large effects on both abiotic and biotic environmental variables, resource mutualisms, in particular, have the potential to influence plant communities. Moreover, the effects of resource mutualists such as nitrogen-fixing rhizobia on diversity and community composition may be more pronounced in nutrient-limited environments. I experimentally manipulated the presence of rhizobia across a nitrogen gradient in early assembling mesocosm communities with identical starting species composition to test how the classic mutualism between nitrogen-fixing rhizobia and their legume host influence diversity and community composition. After harvest, I assessed changes in α-diversity, community composition, β-diversity, and ecosystem properties such as inorganic nitrogen availability and productivity as a result of rhizobia and nitrogen availability. The presence of rhizobia decreased plant community diversity, increased community convergence (reduced β-diversity), altered plant community composition, and increased total community productivity. These community-level effects resulted from rhizobia increasing the competitive dominance of their legume host Chamaecrista fasciculata. Moreover, different non-leguminous species responded both negatively and positively to the presence of rhizobia, indicating that rhizobia are driving both inhibitory and potentially facilitative effects in communities. These findings expand our understanding of plant communities by incorporating the effects of positive symbiotic interactions on plant diversity and composition. In particular, rhizobia that specialize on dominant plants may serve as keystone mutualists in terrestrial plant communities, reducing diversity by more than 40 %.