Unifying the relationships of species richness to productivity and disturbance

Although species richness has been hypothesized to be highest at 'intermediate' levels of disturbance, empirical studies have demonstrated that the disturbance-diversity relationship can be either negative or positive depending on productivity On the other hand, hypothesized productivity diversity relationships can be positive, negative or unimodal, as confirmed by empirical studies. However, it has remained unclear under what conditions each pattern is realized, and there is little agreement about the mechanisms that generate these diverse patterns. In this study, I present a model that synthesizes these separately developed hypotheses and shows that the interactive effects of disturbance and productivity on the competitive outcome of multispecies dynamics can result in these diverse relationships of species richness to disturbance and productivity The predicted productivity diversity relationship is unimodal but the productivity level that maximizes species richness increases with increasing disturbance. Similarly, the predicted disturbance diversity relationship is unimodal but the peak moves to higher disturbance levels with increasing productivity Further, these patterns are well explained by the opposite effects of productivity and disturbance on competitive outcome that are suggested by the change in community composition along these two environmental gradients: higher productivity favours superior competitors while higher disturbance levels favour inferior competitors.     

Neutral communities may lead to decreasing diversity-disturbance relationships: insights from a generic simulation model

Many attempts have been made to confirm or reject the unimodal relationship between disturbance and diversity stated by the intermediate disturbance hypothesis (IDH). However, the reasons why the predictions of the IDH apply or fail in particular systems are not always obvious. Here, we use a spatially explicit, individual-based community model that simulates species coexistence in a landscape subjected to disturbances to compare diversity-disturbance curves of communities with different coexistence mechanisms: neutrality, trade-off mechanism and intraspecific density dependence. We show that the shape of diversity-disturbance curves differs considerably depending on the type of coexistence mechanism assumed: (1) Neutral communities generally show decreasing diversity-disturbance curves with maximum diversity at zero disturbance rates contradicting the IDH, whereas trade-off communities generally show unimodal relationships confirming the IDH and (2) density-dependent mechanisms do increase the diversity of both neutral and trade-off communities. Finally, we discuss how these mechanisms determine diversity in disturbed landscapes.     

Predicting invertebrate diversity from disturbance regimes in forest streams

The link between substrate disturbance and stream invertebrate species richness is often complicated by the fact that substrate disturbance removes both invertebrates and periphyton (a potential food source). It is never clear whether disturbance acts directly on species diversity by removing animals or indirectly by reducing one of their food sources. To examine this relationship invertebrate diversity patterns were examined in 25 forest streams in Urewera National Park, New Zealand, where light attenuation from the forest canopy was postulated to limit periphyton biomass and remove the confounding influence of periphyton on the link between substrate disturbance and invertebrate diversity. Invertebrate species richness declined linearly with increasing substrate disturbance. Although periphyton biomass was comparatively low, species richness was more strongly related to periphyton biomass than to any disturbance measure. The highly mobile nature and terrestrial reproductive stage of many lotic invertebrates suggest that colonisation dynamics may have a more important influence on diversity patterns than monopolisation of resources for population growth. Although both the intermediate disturbance hypothesis and the dynamic equilibrium model encompass colonisation as a critical determinant of diversity both models also require a trade-off between the colonising and competitive ability of individual species; a phenomenon which does not appear to occur widely in lotic communities. Rather, it is postulated that resource levels will set an upper limit to the species richness of a benthic community that can be achieved through colonisation of taxa in the absence of disturbance, while disturbance removes taxa and resets the colonisation process.     

Effects of productivity and disturbance on species richness: a neutral model

Productivity and disturbance are major determinants of species diversity, and results from theoretical models predict that species richness should peak at intermediate levels of both factors. Such "unimodal" responses have been documented in many field and laboratory studies and have usually been attributed to differences among species in competitive ability and/or trade-offs between competitive ability and tolerance to disturbance. Here we show that most documented patterns of disturbance-richness and productivity-richness relationships, as well as the observed interactions between the two factors, can be explained by a simple neutral model where all species are ecologically identical and lack trade-offs in species characteristics. This finding suggests that current neutral theories can be extended to explain patterns of species responses to productivity and disturbance.     

The effects of species pool,dispersal and competition on the diversity–productivity relationship

Aim The diversity–productivity relationship is a controversial issue in ecology. Diversity is sometimes seen to increase with productivity but a unimodal relationship has often been reported. Competitive exclusion was cited initially to account for the decrease of diversity at high productivity. Subsequently, the roles of evolutionary history (species pool size) and dispersal rate have been acknowledged. We explore how the effects of species pool, dispersal and competition combine to produce different diversity–productivity relationships. Methods We use a series of simulations with a spatially explicit, individual‐based model. Following empirical expectations, we used four scenarios to characterize species pool size along the productivity gradient (uniformly low and high, linear increase and unimodal). Similarly, the dispersal rate varied along the productivity gradient (uniformly low and high, and unimodal). We considered both neutral communities and communities with competitive exclusion. Results and main conclusions Our model predicts that competitive interactions will result in unimodal diversity–productivity relationships. The model often predicts unimodal patterns in neutral communities as well, although the decline in richness at high productivity is less than in competing communities. A positive diversity–productivity relationship is simulated for neutral communities when the species pool size increases with productivity and the dispersal rate is high. This scenario is probably more widespread in nature than the others since positive diversity–productivity relationships have been observed more frequently than previously expected, especially in the tropics and for woody species. Our simulated effects of species pool, dispersal and competition on diversity patterns can be linked to empirical observations to uncover mechanisms behind the diversity–productivity relationship.     

Grain size affects the relationship between species richness and above-ground biomass in semi-arid rangelands

ABSTRACT

Background: Discrepancies in the shape of the productivity–diversity relationship may arise from differences in spatial scale. We hypothesised that there is a grain size effect on the productivity–diversity relationship.

Aims: To determine the effect of three sampling grain sizes on the productivity–diversity relationship.

Methods: We applied generalised linear mixed effect models on community data from 735 vegetation plots in the Taleghan rangelands, Iran, sampled at three grain sizes (0.25, 1 and 2 m²) to ascertain plant productivity-diversity patterns, while accounting for the effects of site, plant community type, disturbance, and life form.

Results: Overall, relationships between biomass and plant species richness were unimodal at grain sizes of 0.25 and 1 m², and asymptotical at 2 m². The spurious occurrence of a single large shrub may overwhelm a small-sized sampling unit, resulting in a high estimate of the sample’s biomass relative to species richness. However, the relationship between biomass and species richness at larger grain sizes is more likely to reach an asymptote.

Conclusions: Shrubs are partly responsible for driving the relationship between plant biomass and species richness. Given that the frequency of shrubs is highly variable between small plots but not so in large plots, their presence may result in unimodal productivity–diversity relationships at small but not at large grain sizes.     

The interplay between species' positive and negative interactions shapes the community biomass–species richness relationship

We used an individual-based spatially-explicit model to assess the role of facilitation and plant strategies in shaping the 'community biomass–species richness' relationship. Facilitation had few impacts on community's richness under both the most benign (high community biomass) and the most severe (low community biomass) environments where its intensity was weak. From medium to high environmental severity, facilitation increased community richness, because all plant strategies were facilitated. In contrast, from low to medium environmental severity facilitation decreased community richness, because only the most competitive species were facilitated, which induced a decrease in the richness of the stress-tolerant species overwhelming the increase in richness of the competitive species. Above all, our simulations show how 'strategy-dependent' interactions among species combine to shape the humped-back biomass–species richness relationship. It also demonstrates that facilitative effects might have long-term negative effects on species richness, which result is not included in current facilitation models.     

Productivity-diversity relationships in lake plankton communities

One of the most intriguing environmental gradients connected with variation in diversity is ecosystem productivity. The role of diversity in ecosystems is pivotal, because species richness can be both a cause and a consequence of primary production. However, the mechanisms behind the varying productivity-diversity relationships (PDR) remain poorly understood. Moreover, large-scale studies on PDR across taxa are urgently needed. Here, we examined the relationships between resource supply and phyto-, bacterio-, and zooplankton richness in 100 small boreal lakes. We studied the PDR locally within the drainage systems and regionally across the systems. Second, we studied the relationships between resource availability, species richness, biomass and resource ratio (N:P) in phytoplankton communities using Structural Equation Modeling (SEM) for testing the multivariate hypothesis of PDR. At the local scale, the PDR showed variable patterns ranging from positive linear and unimodal to negative linear relationships for all planktonic groups. At the regional scale, PDRs were significantly linear and positive for phyto- and zooplankton. Phytoplankton richness and the amount of chlorophyll a showed a positive linear relationship indicating that communities consisting of higher number of species were able to produce higher levels of biomass. According to the SEM, phytoplankton biomass was largely related to resource availability, yet there was a pathway via community richness. Finally, we found that species richness at all trophic levels was correlated with several environmental factors, and was also related to richness at the other trophic levels. This study showed that the PDRs in freshwaters show scale-dependency. We also documented that the PDR complies with the multivariate model showing that plant biomass is not mirroring merely the resource availability, but is also influenced by richness. This highlights the need for conserving diversity in order to maintain ecosystem processes in freshwaters.     

Competing drivers lead to non-linear native–exotic relationships in endangered temperate grassy woodlands

Relationships between the diversity and abundance of native versus exotic species underpin management of disturbance regimes for conservation. Theory predicts negative, positive or neutral relationships depending on respective drivers, with greatest potential benefit when natives and exotics show opposing responses to management. We examined drivers of exotic plant cover and relationships with native plant richness using 12-year burning, mowing and grazing experiments in two representative temperate grassy eucalypt woodlands with contrasting histories of frequent versus infrequent disturbance. We hypothesized that disturbance and high resources favour exotics, and assessed whether natives and exotics covary positively due to common external drivers or negatively due to contrasting external drivers and/or competition. Positive relationships with rainfall and disturbance explained >80 % of the variation in exotic cover at both sites, supporting our first hypothesis. Native–exotic relationships were non-linear, with native richness first increasing rapidly with increasing exotic cover, then levelling and beginning to decrease. Common external drivers, particularly inter-annual rainfall, explained initial positive relationships, highlighting a prevalence of positive relationships at long temporal (as well as large spatial) scales. At the historically frequently-burnt site, a concomitant increase in native richness and exotic cover after fire contributed to the positive relationship, indicating a management trade-off. At the long-unburnt site, exotics increased but natives decreased with fire, suggesting dual benefits of low fire frequency. We conclude that relationships between exotic cover and native richness emerge from interactions among external drivers and competitive responses, with responses to external drivers dominating at low resources and negative interactions gaining importance as resources increase.     

Species pools and the "hump-back" model of plant species diversity: an empirical analysis at a relevant spatial scale

We investigated the relative roles of productivity, the species pool, and spatial habitat structure in determining local species richness (alpha diversity) of plant communities within a single, well-defined landscape unit, at spatial and ecological scales where the relationship between community productivity and species diversity often assumes a unimodal or "hump-back" form. At high levels of productivity, the decrease-phase of the unimodal model of the diversity-productivity relationship is typically explained as the dynamic outcome of increased competitive exclusion, but it may also be the passive consequence of a small pool of species possessing attributes necessary to competitively survive in high-fertility environments. We conducted statistical analyses of previously collected data to determine whether variations in local richness in the herbaceous vegetation of a Slovakian mountain valley were best explained by habitat productivity itself (which presumably leads to more intense competition) or by the sizes of the relevant community species pools. We also used measures of spatial habitat structure to investigate the extent to which habitat patchiness influenced patterns of species diversity. In the study system, both community biomass and size of the species pools contributed significantly to local species richness, but the positive effect of the species pools was about twice as important as the negative effect of biomass. The combined area of related associations (alliance area), association perimeter, and habitat patch geometry were all closely related to species pool size.     

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.     

Burial disturbance leads to facilitation among coastal dune plants

There is growing evidence that interactions among plants can be facilitative as well as competitive, but knowledge of how disturbances influence these interactions and how they vary with species diversity is lacking. We manipulated plant density, species diversity (richness), and a burial disturbance in a controlled, complete factorial experiment to test theories about the relationships among species interactions, disturbance, and richness. The hypotheses tested were 1) burial disturbance reduces plant performance at all levels of density and richness, 2) burial disturbance can cause net plant interactions to become more facilitative, and 3) facilitation increases with species richness. Burial decreased plant survival by 60% and biomass by 50%, supporting the hypothesis that burial reduces plant performance. In the control (unburied) treatment, there was no difference in proportion survival or per plant biomass between low and high density plots, meaning that neither competition nor facilitation was detected. In the buried treatment, however, high density plots had significantly greater survival and greater per plant biomass than the low density plots, indicating net facilitative interactions. Thus facilitation occurred in the buried treatment and not in the unburied control plots, supporting the hypothesis that facilitation increases with increasing disturbance severity. The hypothesis that facilitation increases with increasing species richness was not supported. Richness did not affect survival or biomass, and there was no richness by burial treatment interaction, indicating that richness did not influence the response of the community to burial. The influence of the disturbance on plant interactions was thus consistent across levels of richness, increasing the generality of the relationship between disturbance and facilitation.     

The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Climate-induced temporal variation in the productivity–diversity relationship

Spatial scales are known to influence the form of the productivity–diversity relationship, but less attention has been given to the influence of temporal scales. Interannual climatic variation in semi-arid ponderosa pine–bunchgrass ecosystems causes significant year-to-year differences in herbaceous production. We hypothesized that unimodal (or 'hump-backed') relationships would be detected between herbaceous production and species richness in wet years, whereas positive logarithmic relationships would be detected in dry years. We analyzed nine years of herbaceous production and species richness data and used Akaike's information criterion (AICc) to determine the weight of evidence for each model in each year. As predicted, species richness exhibited a unimodal relationship to herbaceous production in wet years; however, richness exhibited a logarithmic relationship with herbaceous production in dry years. These results suggest that competitive exclusion occurred within this semi-arid plant community in years of high production when enough moisture was available to drive abundant plant growth. Thus, just as it is important to sample broad spatial variation in production to detect the full unimodal productivity–diversity relationship, it is also important to recognize that the full unimodal curve may be undetectable in less productive dry years in semi-arid ecosystems.     

Experimental test of the intermediate disturbance hypothesis: frequency effects of emersion on fouling communities

The validity of predictions derived from the intermediate disturbance hypothesis (IDH) was tested in situ by manipulating mussel dominated Western Baltic fouling communities. Assemblages of two different successional stages, 3 and 12 months old, underwent a 3-month period of disturbance treatment in terms of various frequencies of emersion. Emersion frequency levels ranged from 1×15 to 48×15 min emersion day⁻¹. The study on the 3-month-old communities was repeated in 2 subsequent study years. Species richness, evenness and diversity (Shannon index) were recorded to measure the effects of frequency treatments on community structure.The IDH was confirmed in the first year, when diversity was found to be a unimodal function of the applied emersion frequency gradient. Diversity-disturbance relationships were inverse unimodal or non-significant in the second year, which was true for both successional stages. This ambiguous picture partially confirms the validity of the mechanisms proposed by the IDH, but also shows that their forcing can be masked by fluctuations in environmental parameters, such as climatic conditions. Diversity increased again under severe disturbance conditions, due to a disturbance-induced change in community structure, namely the shift from mussel to algal dominance. This is a new aspect in the discussion concerning disturbance-diversity relationships.     

Consistent Richness-Biomass Relationship across Environmental Gradients in a Marine Macroalgal-Dominated Subtidal Community on the Western Antarctic Peninsula

Biodiversity loss has spurred the biodiversity-ecosystem functioning research over a range of ecosystems. In Antarctica, however, the relationship of taxonomic and functional diversity with ecosystem properties (e.g., community biomass) has received less attention, despite the presence of sharp and dynamic environmental stress gradients that might modulate these properties. Here, we investigated whether the richness-biomass relationship in macrobenthic subtidal communities is still apparent after accounting for environmental stress gradients in Fildes Bay, King George Island, Antarctica. Measurements of biomass of mobile and sessile macrobenthic taxa were conducted in the austral summer 2013/4 across two environmental stress gradients: distance from nearest glaciers and subtidal depth (from 5 to 30 m). In general, community biomass increased with distance from glaciers and water depth. However, generalised additive models showed that distance from glaciers and depth accounted for negligible proportions of variation in the number of functional groups (i.e., functional richness) and community biomass when compared to taxonomic richness. Functional richness and community biomass were positive and saturating functions of taxonomic richness. Large endemic, canopy-forming brown algae of the order Desmarestiales dominated the community biomass across both gradients. Accordingly, differences in the composition of taxa accounted for a significant and large proportion (51%) of variation in community biomass in comparison with functional richness (10%). Our results suggest that the environmental factors here analysed may be less important than biodiversity in shaping mesoscale (several km) biomass patterns in this Antarctic system. We suggest that further manipulative, hypothesis-driven research should address the role of biodiversity and species’ functional traits in the responses of Antarctic subtidal communities to environmental variation.     

Seasonal changes in the relationship between plant species richness and community biomass in early succession

We analysed the relationship between plant species richness and productivity on first-year-old fields at two similar sites in central Europe. At both sites, a wide range of productivity levels was available resulting from different long-term fertilisation. In order to identify underlying mechanisms of the species richness–productivity relationship we included the seasonal dynamics and the number of individuals of each species in our analysis. We sampled 10 and 21 plots, respectively, at the two sites in May, June and July by harvesting all aboveground parts of vascular plants in 0.25 m² subplots. Species richness, number of individuals of each species and community biomass as a surrogate of productivity were recorded in each sample.At one site, the relationship between species richness and biomass was significantly positive in the May and June harvest. This relationship disappeared in the July harvest due to a reduction in species richness at high productivity levels. The relations between species richness and number of individuals and between number of individuals and biomass paralleled the species richness–productivity relation but the individual number–biomass relationship remained positive until the last harvest. Between-species differences in individual number–community biomass relationships and their seasonal dynamics revealed “interspecific competitive exclusion” even though the species richness–biomass relationships were not negative or hump-shaped. At the second site, species richness was not related to productivity or to number of individuals. Our study demonstrated the importance of temporal dynamics and regional processes in understanding species richness–productivity patterns.     

Patterns in Species Persistence and Biomass Production in Soil Microcosms Recovering from a Disturbance Reject a Neutral Hypothesis for Bacterial Community Assembly

The neutral theory of biodiversity has emerged as a major null hypothesis in community ecology. The neutral theory may sufficiently well explain the structuring of microbial communities as the extremely high microbial diversity has led to an expectation of high ecological equivalence among species. To address this possibility, we worked with microcosms of two soils; the microcosms were either exposed, or not, to a dilution disturbance which reduces community sizes and removes some very rare species. After incubation for recovery, changes in bacterial species composition in microcosms compared with the source soils were assessed by pyrosequencing of bacterial 16S rRNA genes. Our assays could detect species with a proportional abundance ≥ 0.0001 in each community, and changes in the abundances of these species should have occurred during the recovery growth, but not be caused by the disturbance per se. The undisturbed microcosms showed slight changes in bacterial species diversity and composition, with a small number of initially low-abundance species going extinct. In microcosms recovering from the disturbance, however, species diversity decreased dramatically (by > 50%); and in most cases there was not a positive relationship between species initial abundance and their chance of persistence. Furthermore, a positive relationship between species richness and community biomass was observed in microcosms of one soil, but not in those of the other soil. The results are not consistent with a neutral hypothesis that predicts a positive abundance-persistence relationship and a null effect of diversity on ecosystem functioning. Adaptation mechanisms, in particular those associated with species interactions including facilitation and predation, may provide better explanations.     

Competitive exclusion,beta diversity,and deterministic vs. stochastic drivers of community assembly

Species diversity has two components – number of species and spatial turnover in species composition (beta‐diversity). Using a field experiment focusing on a system of Mediterranean grasslands, we show that interspecific competition may influence the two components in the same direction or in opposite directions, depending on whether competitive exclusions are deterministic or stochastic. Deterministic exclusions reduce both patch‐scale richness and beta‐diversity, thereby homogenising the community. Stochastic extinctions reduce richness at the patch scale, but increase the differences in species composition among patches. These results indicate that studies of competitive effects on beta diversity may help to distinguish between deterministic and stochastic components of competitive exclusion. Such distinction is crucial for understanding the causal relationship between competition and species diversity, one of the oldest and most fundamental questions in ecology.     

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.