Extending the quasi-neutral concept

In caricature, the equilibrium paradigm of community ecology states that plant communities are stable entities consisting of competing species - and that such species coexist because each has its “niche”. This paradigm, in its extreme, has been dead for some time. Nevertheless, it has yet to be replaced by a credible “non” — equilibrium paradigm. The quasi-neutral concept of plant communities, proposed by Kristjan Zobel,Folia Geobot. 36: 3–8, 2001, possesses some of the key ingredients of a nonequilibrium theory of diversity. It recognizes that there are inescapable relationships between diversity at different scales, that similarity can influence the rate of competitive exclusion, that successional change is typically associated with changes in life form, and that rarefaction (i.e. “sampling artifacts”) can have strong effects on fine-scale diversity. However, the current formulation of the quasi-neutral concept is incomplete in that it relies on an unrealistic definition of community, it assumes that random sampling means that species richness at one scale will be linearly related to richness at finer scales, it ignores the possibility of fine-scale processes producing broader-scale patterns, and it avoids the subject of fine-scale environmental heterogeneity. But the most serious limitation of the quasi-neutral concept is that similarity of species alone is not sufficient to allow indefinite coexistence. I present the results of a simple simulation to demonstrate that: (1) identical species will eventually be lost from communities due to stochastic “drift”, (2) slight variations in reproductive rates accelerate this loss, but (3) adding a miniscule “cost of commonness” to the model allows the indefinite coexistence of species. I conclude that the quasi-neutral model cannot work without some kind of trade-off.     

Species richness peaks for intermediate levels of biomass in a fractal succession with quasi-neutral interactions

The mechanisms that promote species richness, including net community interactions, are considered central to the investigation of the consequences of biodiversity loss for ecosystem functioning. Recently, some empirical studies at large spatiotemporal scales suggest that increasing species richness within natural communities results in a finer division of biomass among species rather than an increase in total biomass. In parallel, the most common pattern observed in nature is the peaked relationship between diversity and productivity estimated as total biomass. Thus, the aim of our study is to provide model predictions for the diversity–biomass relationship with various levels of net species interactions within communities: negative, neutral, quasi-neutral and positive. Using a scaling relationship between the number of species and total community biomass, we propose a new self-similar process of biomass partitioning during a community assembly process. At each step of the succession, K more species appear that are A times less abundant on average giving K=A^d; the parameter d being a fractal dimension related to the nature of interactions among coexisting species. Our results, compared to those from meta-analyses about empirical diversity–productivity relationships, illustrate that quasi-neutral interactions among coexisting species lead to the most commonly observed pattern: an 'envelope' where diversity peaks at intermediate values of total biomass, i.e. that the area below the hump-backed line (considered as the upper boundary) is filled with data points.     

On the species-pool hypothesis and on the quasi-neutral concept of plant community diversity

5.1. According to the results from studying a broad variety of Estonian herbaceous communities (4.11) the question in 1.14 should be answered as:selection from a regional species pool into an actual species pool and selection from actual species pool into a microsite are mostly random and neutral processes and they are not directed significantly by interspecific competition. 5.2. Yet, the formation of a diversity pattern should be called aquasi-neutral process,mainly because the exclusion of species from communities due to asymmetric light competition is common during succession (when taller species outcompete shorter ones).     

Genetic and morphological patterns show variation in frequency of hybrids between Ipomopsis (Polemoniaceae) zones of sympatry

Variation in rates of hybridization among zones of sympatry between a pair of species provides a useful window into the effect of local conditions on the evolution of reproductive isolation. We employed floral morphological traits and neutral genetic markers to quantify the frequency of individuals intermediate to the two parental species in two zones of sympatry between Ipomopsis aggregata and I. tenuituba, using clustering methods that make no a priori assumptions about population structure. The sites differed not only in the frequency of intermediate individuals, but also in climate, pollinator abundance and behavior and spatial structure of plant populations. Both floral traits, which are likely to be under natural selection and molecular markers, which are quasi-neutral, indicated more population structure at one site than the other, the pattern being more pronounced for floral morphology. One likely explanation for this difference between sites is that local ecological conditions, particularly pollinator choice of flowers, have promoted different rates of hybridization between these species. Hence, the evolution of reproductive isolation might depend in part on local conditions, and thus differ among populations of the same pair of species.     

DNA variants of the MHC show location-specific convergence between sheep, goat and cattle

Interspecific convergent evolution in sheep, goat and cattle was analysed with the help of orthologous microsatellite markers. Six of the loci are located in the major histocompatibility complex (MHC) region and three on different chromosomes. Samples from at least 60 animals per autochthonous breed of the three species were collected in central and southeast Anatolia (Turkey) as well as Baden-Württemberg (Germany). Allelic diversity, heterozygosity, population differentiation and genetic distances were calculated. The loci were polymorphic in all species and breeds. Apart from MSDRB, the loci linked to the MHC were similarly polymorphic as compared to the other loci. Allele numbers in the Turkish sheep and in the cattle breeds were higher than in the other breeds. The predominant occurrence of distinct allele lengths per locus differed depending on the species. For the three geographic locations, the genetic distances between species based on the MHC loci were significantly closer in comparison with distances based on quasi-neutral loci. This indicates convergent evolution of the MHC loci between sheep, goat and cattle caused by effects of location and demonstrates an approach for quantifying influences of adaptation on genetic variability.     

Local richness-species pool ratio: A consequence of the species-area relationship

A constant ratio between species richness estimated at the local and regional scale is interpreted as a proof of quasi-neutral unsaturated communities. Based on Zobel’s model of plant community (Zobel,Folia Geobot. 36: 3–8, 2001) we tested the methodology of the species-pool concept by comparing the saturated and unsaturated communities generated by spatially-explicit mechanistic simulations with known assembly rules. Tests show that local-regional species plots can be applied to distinguish saturated vs. unsaturated communities, however, the outcome of tests, i.e. the relationship between local and regional richness depends on the size of the areas compared. Independently from the mechanisms controlling diversity, trivial saturation will appear if one of the scales is either too small or too broad because species-area curves are bound at these extreme scales. Similarly, trivial unsaturaton will appear if the two scales compared are close to each other. The application of species-area curves is useful because they help to find scales for non-trivial relationships. Field tests reporting quasi-neutrality and unsaturated plant communities were performed at the intermediate scales of the corresponding species-area curves, and they were estimated from heterogeneous samples. Therefore, this field evidence might be biased by scaling artefacts. We propose to reanalyze the field evidence with solid scaling conventions and to restrict the concept of quasi-neutrality to subordinated functional groups based on the following hypotheses: (1) neutrality will appear within subordinated guilds as a consequence of the hierarchical structure of plant communities; (2) the lower a guild in the hierarchy the higher neutrality of within-layer processes detected; (3) quasi-neutrality found at the community level is not a proof of community-level neutrality but it is due to the higher number of subordinated species in the samples.     

Functional redundancy in ecology and conservation

Multiple studies have shown that biodiversity loss can impair ecosystem processes, providing a sound basis for the general application of a precautionary approach to managing biodiversity. However, mechanistic details of species loss effects and the generality of impacts across ecosystem types are poorly understood. The functional niche is a useful conceptual tool for understanding redundancy, where the functional niche is defined as the area occupied by a species in an n-dimensional functional space. Experiments to assess redundancy based on a single functional attribute are biased towards finding redundancy, because species are more likely to have non-overlapping functional niches in a multi-dimensional functional space. The effect of species loss in any particular ecosystem will depend on i) the range of function and diversity of species within a functional group, ii) the relative partitioning of variance in functional space between and within functional groups, and iii) the potential for functional compensation (degree of functional niche overlap) of the species within a functional group. Future research on functional impairment with species loss should focus on identifying which species, functional groups, and ecosystems are most vulnerable to functional impairment from species loss, so that these can be prioritized for management activities directed at maintaining ecosystem function. This will require a better understanding of how the organization of diversity into discrete functional groups differs between different communities and ecosystems.     

The importance of rare species: a trait‐based assessment of rare species contributions to functional diversity and possible ecosystem function in tall‐grass prairies

The majority of species in ecosystems are rare, but the ecosystem consequences of losing rare species are poorly known. To understand how rare species may influence ecosystem functioning, this study quantifies the contribution of species based on their relative level of rarity to community functional diversity using a trait‐based approach. Given that rarity can be defined in several different ways, we use four different definitions of rarity: abundance (mean and maximum), geographic range, and habitat specificity. We find that rarer species contribute to functional diversity when rarity is defined by maximum abundance, geographic range, and habitat specificity. However, rarer species are functionally redundant when rarity is defined by mean abundance. Furthermore, when using abundance‐weighted analyses, we find that rare species typically contribute significantly less to functional diversity than common species due to their low abundances. These results suggest that rare species have the potential to play an important role in ecosystem functioning, either by offering novel contributions to functional diversity or via functional redundancy depending on how rare species are defined. Yet, these contributions are likely to be greatest if the abundance of rare species increases due to environmental change. We argue that given the paucity of data on rare species, understanding the contribution of rare species to community functional diversity is an important first step to understanding the potential role of rare species in ecosystem functioning.     

南亚热带森林常见种和偶见种的功能多样性特征

群落特征可能取决于物种在功能实体功能特征的独特组合（FE）中的分布。了解这些分布是保护生物多样性和维持生态系统功能的基础。鼎湖山南亚热带常绿阔叶林群落物种丰富,根据物种的多度来区分常见种（每hm²超过一个个体）和偶见种（每hm²少于一个个体）。基于功能实体,分别计算功能实体数量、功能冗余度、功能脆弱度和功能过度冗余度来描述每个样方中常见种和偶见种的功能多样性。采用Wilcoxon检验检测常见种和偶见种功能性状的差异,通过线性回归分析常见种和偶见种的功能多样性与物种数量的关系,采用冗余分析（RDA）影响常见种和偶见种的功能多样性的环境因子。研究结果表明：（1）鼎湖山南亚热带常绿阔叶林群落中功能性状在常见种和偶见种间差异显著。（2）群落中数量较低的偶见种保持着较高比例的功能实体数量。功能实体数量、功能冗余度与常见种和偶见种数量显著正相关,功能过度冗余度与偶见种数量显著正相关,功能脆弱度与常见种和偶见种的物种数量呈显著负相关。（3）海拔是影响常见种功能多样性最关键的地形因子,凹凸度是影响偶见种功能多样性最关键的地形因子。土壤含水量是影响常见种功能多样性最关键的土壤因子,速效钾是影响偶见种功能多样性最关键的环境因子。研究结果表明,尽管物种丰富的热带森林具有较高的功能冗余,但其提供的保险效应不能抵消生态系统的功能脆弱性。减少具有独特功能实体和偶见种的损失是避免热带森林生态系统功能损失的有效和必要的方法。     

Co-occurrence of tree species at fine spatial scale in a woodland cerrado,southeastern Brazil

Species co-occurrence at fine spatial scales is expected to be nonrandom in relation to species phylogenetic relatedness and functional similarity. On the one hand, closely related species that occur together and experience similar environmental conditions are likely to share phenotypic traits due to the process of environmental filtering. On the other hand, species that are too similar are unlikely to co-occur due to competitive exclusion. We surveyed a woodland cerrado, southeastern Brazil, to test whether co-occurrence in tree species shows functional or phylogenetic structuring at fine spatial scale. Searching for correlations between an index of species co-occurrence and both functional trait differences and phylogenetic distances, we provided evidence for a predominant role of environment filters in determining the co-occurrence of functionally similar tree species in cerrado. However, we did not find any effect of phylogenetic relatedness on tree species co-occurrence. We suggest that the phylogenetic relatedness of co-occurring cerrado tree species did not present a pattern, because the species functional traits were randomly distributed on the phylogeny. Thus, phylogenetic relatedness and functional similarity do not seem to limit the co-occurrence at fine spatial scale of cerrado tree species.     

Distance-Based Functional Diversity Measures and Their Decomposition: A Framework Based on Hill Numbers

Hill numbers (or the “effective number of species”) are increasingly used to characterize species diversity of an assemblage. This work extends Hill numbers to incorporate species pairwise functional distances calculated from species traits. We derive a parametric class of functional Hill numbers, which quantify “the effective number of equally abundant and (functionally) equally distinct species” in an assemblage. We also propose a class of mean functional diversity (per species), which quantifies the effective sum of functional distances between a fixed species to all other species. The product of the functional Hill number and the mean functional diversity thus quantifies the (total) functional diversity, i.e., the effective total distance between species of the assemblage. The three measures (functional Hill numbers, mean functional diversity and total functional diversity) quantify different aspects of species trait space, and all are based on species abundance and species pairwise functional distances. When all species are equally distinct, our functional Hill numbers reduce to ordinary Hill numbers. When species abundances are not considered or species are equally abundant, our total functional diversity reduces to the sum of all pairwise distances between species of an assemblage. The functional Hill numbers and the mean functional diversity both satisfy a replication principle, implying the total functional diversity satisfies a quadratic replication principle. When there are multiple assemblages defined by the investigator, each of the three measures of the pooled assemblage (gamma) can be multiplicatively decomposed into alpha and beta components, and the two components are independent. The resulting beta component measures pure functional differentiation among assemblages and can be further transformed to obtain several classes of normalized functional similarity (or differentiation) measures, including N-assemblage functional generalizations of the classic Jaccard, Sørensen, Horn and Morisita-Horn similarity indices. The proposed measures are applied to artificial and real data for illustration.     

Taxonomic and functional approaches reveal different responses of ant assemblages to land-use changes

Land-use change is well documented to cause species loss. However, our understanding of the effects of land-use change on other aspects of biodiversity is still limited. We evaluated if different land-use changes (Eucalyptus plantation and planted pasture) affect ant species and functional groups in similar ways across three Cerrado vegetation types (grassland, savanna and savanna-forest). We found that ant species and functional responses differed with land-use change in relation to frequency of occurrence and habitat specificity and fidelity. Land-use change affected species frequency of occurrence but not functional groups, indicating that species are more sensitive than functional groups to habitat transformation. Native habitats had different indicator species and functional groups compared with converted habitats. However, we did not find functional group indicators of converted habitats in any vegetation type; indicating that there is no specificity and fidelity of functional group to converted habitats and that such an approach is less sensitive to land-use changes. In savanna and savanna-forest, species and functional groups showed the same response in relation to composition with differences between native and converted habitats. Thus, functional groups will be lost when ant species are lost. In grasslands, functional group composition was similar between native and converted habitats indicating a turnover of species within functional groups. We demonstrate that both Eucalyptus plantation and planted pasture affect ant species and functional groups in different ways, with negative impacts both taxonomically and less so functionally. Therefore, we show that the two aspects of biodiversity can respond independently to land-use changes and, hence, the importance of using both taxonomic and functional group approaches to evaluate the effects of land-use change on biodiversity in savanna systems.     

The consequences of fluctuating selection for isozyme polymorphisms in Daphnia

Temporal sequences of allele frequencies in natural populations of Daphnia are analyzed to obtain the mean and variance of the selection coefficient for both asexual and sexual phases. In general, the alleles at enzyme loci appear to be quasi-neutral. Although significant variation exists for the estimated selection coefficients, the means are in all cases close to zero. Estimates of the variance of selection intensity are applied to existing models to demonstrate the implications of fluctuating selection for the spatial and temporal distribution of gene frequencies in Daphnia. The empirical and analytical results are shown to provide a possible solution to some previously puzzling aspects of Daphnia population genetic surveys. Neither genetic drift nor diversifying selection are necessary conditions for the local diversification of gene frequencies.     

Evolution of a complex phenotype with biphasic ontogeny: Contribution of development versus function and climatic variation to skull modularity in toads

The theory of morphological integration and modularity predicts that if functional correlations among traits are relevant to mean population fitness, the genetic basis of development will be molded by stabilizing selection to match functional patterns. Yet, how much functional interactions actually shape the fitness landscape is still an open question. We used the anuran skull as a model of a complex phenotype for which we can separate developmental and functional modularity. We hypothesized that functional modularity associated to functional demands of the adult skull would overcome developmental modularity associated to bone origin at the larval phase because metamorphosis would erase the developmental signal. We tested this hypothesis in toad species of the Rhinella granulosa complex using species phenotypic correlation pattern (P‐matrices). Given that the toad species are distributed in very distinct habitats and the skull has important functions related to climatic conditions, we also hypothesized that differences in skull trait covariance pattern are associated to differences in climatic variables among species. Functional and hormonal‐regulated modules are more conspicuous than developmental modules only when size variation is retained on species P‐matrices. Without size variation, there is a clear modularity signal of developmental units, but most species have the functional model as the best supported by empirical data without allometric size variation. Closely related toad species have more similar climatic niches and P‐matrices than distantly related species, suggesting phylogenetic niche conservatism. We infer that the modularity signal due to embryonic origin of bones, which happens early in ontogeny, is blurred by the process of growth that occurs later in ontogeny. We suggest that the species differing in the preferred modularity model have different demands on the orbital functional unit and that species contrasting in climate are subjected to divergent patterns of natural selection associated to neurocranial allometry and T3 hormone regulation.     

The worldwide impact of urbanisation on avian functional diversity

Urbanisation is driving rapid declines in species richness and abundance worldwide, but the general implications for ecosystem function and services remain poorly understood. Here, we integrate global data on bird communities with comprehensive information on traits associated with ecological processes to show that assemblages in highly urbanised environments have substantially different functional composition and 20% less functional diversity on average than surrounding natural habitats. These changes occur without significant decreases in functional dissimilarity between species; instead, they are caused by a decrease in species richness and abundance evenness, leading to declines in functional redundancy. The reconfiguration and decline of native functional diversity in cities are not compensated by the presence of exotic species but are less severe under moderate levels of urbanisation. Thus, urbanisation has substantial negative impacts on functional diversity, potentially resulting in impaired provision of ecosystem services, but these impacts can be reduced by less intensive urbanisation practices.     

Hollow oaks and beetle functional diversity: Significance of surroundings extends beyond taxonomy

Veteran hollow oaks (Quercus spp.) are keystone structures hosting high insect diversity but are declining in numbers due to intensification of land use and the abandonment of traditional management. The loss of this vital habitat is resulting in a reduction of biodiversity, and this likely has consequences for ecosystem functioning, especially if functional diversity is reduced. A considerable amount of research has been done on predictors of beetle taxonomic diversity in veteran oaks, but predictors of functional diversity have remained largely unexplored. The aim of this study was to establish whether the features and surroundings of veteran oaks are related to functional diversity within three functional groups of beetles (decomposers, predators, and flower visitors) and determine whether species richness and functional diversity within the groups are dependent on the same predictors. Sampling was carried out intermittently between 2004 and 2011 on 61 veteran oaks in Southern Norway. Of the 876 beetle species that were collected, 359 were determined to be decomposers, 284 were predators, and 85 were flower visitors. Species richness and functional diversity in all groups were consistently higher in traps mounted on veteran oaks in forests than in open landscapes. However, additional predictors differed between groups, and for species richness and functional diversity. Decomposer species richness responded to tree vitality, while functional diversity responded to habitat connectivity, predator species richness responded to regrowth of shrubs while functional diversity responded to tree circumference, and flower visitor richness and functional diversity did not respond to any additional predictors. Previous studies have found that the features and surroundings of veteran oaks are important for conservation of taxonomic diversity, and the results from this study indicate that they are also important for functional diversity within multiple functional groups.     

More species,but all do the same: contrasting effects of flood disturbance on ground beetle functional and species diversity

The role of habitat disturbance on biodiversity is central as it promotes changes in ecological systems. That said, still little is known about the functional consequences of such changes. Functional diversity can be used to revealing more mechanistically the disturbance effects on communities by considering the richness and the distribution of traits among the species. Here we analyzed the response of functional and species diversity of ground beetles to flood disturbance to better understand the functioning of alluvial invertebrate communities. Ground beetles were sampled in periodically flooded grasslands along the Elbe River in Germany. We used generalized linear mixed effects models to unveil the relationships between flood disturbance, species and functional diversity, respectively. We measured different components of functional diversity (functional richness, evenness, dispersion, and divergence) and analyzed species diversity by means of rarefied species richness, abundances, evenness and Simpson's diversity. We found contrasting relationships in that most species diversity measures peaked at highest disturbance levels, while most functional diversity measures decreased with increasing disturbance intensities. Inversed relationships between species and functional diversity are rarely observed, as most studies report on positive correlations. We explain increasing species diversity with a higher amount of resources available in highly disturbed sites. Decreasing functional diversity is best explained through the convergence of species traits by flood disturbance and uneven resource exploitation in highly disturbed plots (low functional evenness), suggesting strong impacts from functionally different generalist species in floodchannels. We show that the amount of resources available, and how these resources are exploited, play major roles in the functioning of floodplain ground beetle communities.     

高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

Does functional redundancy of communities provide insurance against human disturbances? An analysis using regional-scale stream invertebrate data

Human-induced reductions in species richness might alter the quality of ecosystem services when the remaining species are not able to substitute the functions provided by extirpated species. We examined how human disturbances (nutrient enrichment, land use intensification, instream habitat degradation and the presence of alien species) influence the species richness of stream invertebrates. Stream invertebrates (425 native species) were collected by kick and sweep sampling technique at 274 stream sites covering the entire area of Hungary. We measured the species richness, functional richness (i.e. number of unique functional roles provided by community members) and functional redundancy (i.e. the functional insurance of the community) using information on the feeding habits of each species. To remove the effect of natural variability, we tested the effect of stressors on the residuals of models relating species richness, functional richness and functional redundancy with natural environmental gradients. Our results showed that species richness was negatively influenced by instream habitat degradation and nutrient enrichment. Independent of the way of quantifying functional richness and functional redundancy, we found that functional richness is more sensitive to human impact than functional redundancy of stream invertebrates. The finding that a reduction of species richness is associated with a loss of unique functional roles (functional richness) is important for conservation issues, because the number of unique functional roles is usually regarded as driver of ecosystem functioning.