Using rarity to infer how dendritic network structure shapes biodiversity in riverine communities

Dispersal of organisms connects physical localities, but the strength of connection varies widely. Variability in the influence of dispersal can be predictable in sharply defined networks like river systems because some sections of the network are more isolated, leading to different balances of local (i.e. environmental filtering, species interactions) and regional (i.e. dispersal‐driven) processes in structuring communities. We examined the influence of spatial isolation on the relative contributions of α‐ and β‐diversity to regional (γ) diversity, and examined how that influence differed between common and rare species in stream macroinvertebrate communities. One explanation for rarity on a regional scale is that common species are habitat generalists while rare species are specialists. Therefore, common species should be influenced more by dispersal‐driven processes while rare species should be more influenced by local processes. We predicted that for rare taxa, β‐diversity should represent a higher fraction of γ‐diversity in isolated headwaters but that differences between rare and common taxa with regard to the contribution of β‐diversity to γ‐diversity should be less distinct in well‐connected mainstem habitats. To test these predictions, we used macroinvertebrate communities from 634 sites across 22 watersheds. Regardless of rarity, β‐ and γ‐diversity were higher in headwaters compared to mainstems. However, α‐diversity was similar regardless of isolation for rare assemblages. But contrary to our predictions, common assemblages of predators and herbivores did exhibit differences in α‐diversity between locations. Our predictions were strongly supported for two guilds of consumers, the detritivores and collectors, but less so for herbivores and predators. However, these results make sense considering differences in life histories between the groups. For detritivores and collectors, species turnover (β‐diversity) was higher in isolated regions in river networks, and rarity exacerbated this effect, resulting in higher regional diversity of rare species, supporting the general theory that rarity reflects habitat specialization.     

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.     

Variations in species and functional plant diversity along climatic and grazing gradients

Different components of biodiversity may vary independently of each other along environmental gradients giving insights into the mechanisms that regulate species coexistence. In particular, the functional diversity (FD) or the presence of rare or endemic species in natural assemblages do not necessarily increase with species diversity. We studied if different components of plant species diversity (species richness, Simpson diversity, evenness) varied similarly to FD (measured as a generalization of the Simpson index) and rarity along grazing intensity and climatic gradients. Plots under different sheep grazing regimes (high and low intensity, abandonment) were surveyed in five locations along a climatic gradient in north-eastern Spain, from semi-arid lowland to moist upland locations. Variation in species diversity, functional diversity and rarity followed different patterns. Species diversity was lowest in water-stressed environments (arid locations and southern aspects) and increased with grazing more makedly in humid locations. The FD was comparable between the most species-poor and species-rich locations and decreased with grazing in the moistest location, i.e. where species diversity markedly increased. The FD did not show a strong correlation with species richness nor with the Simpson index and less specious communities could show the highest functional diversity. The rarest species in the region were more frequently found in the abandoned areas, which held the lowest species diversity. Consequently, the mechanisms that enhance the diversity of species do not necessarily support a functional differentiation among those species or the maintenance of rare species in a region. We hypothesize that the degree of dependence of functional diversity on species diversity might be mostly related to the amplitude of the species' traits pool and on how species partition the niche space available.     

Both rare and common species make unique contributions to functional diversity in an ecosystem unaffected by human activities

Aim

Rare species typically contribute more to functional diversity than common species. However, humans have altered the occupancy and abundance patterns of many species—the basis upon which we define “rarity.” Here, we use a globally unique dataset from hydrothermal vents—an untouched ecosystem—to test whether rare species over‐contribute to functional diversity.

Location

Juan de Fuca Ridge hydrothermal vent fields, Northeast Pacific Ocean.

Methods

We first conduct a comprehensive review to set up expectations for the relative contributions of rare and common species to functional diversity. We then quantify the rarity and commonness of 37 vent species with relevant trait information to assess the relationship between rarity and functional distinctiveness—a measure of the uniqueness of the traits of a species relative to traits of coexisting species. Next, we randomly assemble communities to test whether rare species over‐contribute to functional diversity in artificial assemblages ranging in species richness. Then, we test whether biotic interactions influence functional diversity contributions by comparing the observed contribution of each species to a null expectation. Finally, we identify traits driving functional distinctiveness using a distance‐based redundancy analysis.

Results

Across functional diversity metrics and species richness levels, we find that both rare and common species can contribute functional uniqueness. Some species always offer unique trait combinations, and these species host bacterial symbionts and provide habitat complexity. Moreover, we find that contributions of species to functional diversity may be influenced by biotic interactions.

Main conclusions

Our findings show that many common species make persistent, unique contributions to functional diversity. Thus, it is key to consider whether the abundance and occupancy of species have been reduced, relative to historical baselines, when interpreting the contributions of rare species to functional diversity. Our work highlights the importance of testing ecological theory in ecosystems unaffected by human activities for the conservation of biodiversity.     

Annual variation of community biomass is lower in more diverse stream fish communities

Anthropogenic influences have disproportionally affected freshwater ecosystems, and a loss of biodiversity is forecasted to greatly reduce ecosystem function and services. Loss of species may destabilize communities by limiting the stabilizing forces of compensatory dynamics and/or statistical averaging, both of which are effects that can buffer variation in aggregate community properties. Currently, support for positive diversity‐stability relationships stems from experiments with simple communities at small spatial and temporal scales, and application to natural communities is limited. Using a long‐term dataset of 35 stream fish communities matched with hydrologic data, we show that community stability (annual variation of standing biomass of fishes) was less variable in more species‐rich communities and was not associated with stream hydrology. Only the statistical averaging model of community stability was consistent with observed patterns of lower biomass variation in more species‐rich communities. Our findings suggest anthropogenically induced extirpation of vertebrate consumers may lower community biomass stability in complex ecosystems.     

海南岛坝王岭热带天然要植物物种多样性研究

采用植物群落学和植物种群学的研究方法,从数量和质量 特征两个方面研究了海南岛坝王岭热带天然林的植物物种多样性。结果表明：⑴该地区不同植物群落的优势种类组成差别较大,彼此之间的物种相似性水平低;⑵相对于低地雨林而言,该地区的山地雨林物种数量较多,物种均匀度较高,主要优势种的优势程度较弱;⑶在优先考虑物种稀有性的前提下,根据各物物种在群落中所处的位置及其分布情况,可将该地区的植物物种分为珍稀濒危种、特     

You don’t belong here: explaining the excess of rare species in terms of habitat,space and time

Ecological communities are composed of a few common and several rare species. Many studies have evaluated the shape of abundance distribution curves, but few studies have assessed the causes of rarity. Using a dataset of stream macroinvertebrates, we investigated whether the excess of rare species in three focal communities of stones in riffles were common 1) in other habitats at the same stream site and period of sampling (environment), 2) in other stream sites in the same habitat and period of sampling (space), and 3) in other years in the same stream site and habitat (time). We observed that around 28% of the rare species were common in other habitats (environment), stream sites (space) or years (time). Among the three factors, rarity was mostly explained by habitat type, whereas a significant portion of the rare species in riffles were common in pools, submerged roots of terrestrial plants or in partially submerged moss patches. This result suggests that the presence in non‐optimum habitat is a strong determinant of the rarity observed in natural communities and most rare species are due to sampling artifacts or accidentally sampled transient species.     

Understanding the Spatio-Temporal Response of Coral Reef Fish Communities to Natural Disturbances: Insights from Beta-Diversity Decomposition

Understanding how communities respond to natural disturbances is fundamental to assess the mechanisms of ecosystem resistance and resilience. However, ecosystem responses to natural disturbances are rarely monitored both through space and time, while the factors promoting ecosystem stability act at various temporal and spatial scales. Hence, assessing both the spatial and temporal variations in species composition is important to comprehensively explore the effects of natural disturbances. Here, we suggest a framework to better scrutinize the mechanisms underlying community responses to disturbances through both time and space. Our analytical approach is based on beta diversity decomposition into two components, replacement and biomass difference. We illustrate this approach using a 9-year monitoring of coral reef fish communities off Moorea Island (French Polynesia), which encompassed two severe natural disturbances: a crown-of-thorns starfish outbreak and a hurricane. These disturbances triggered a fast logistic decline in coral cover, which suffered a 90% decrease on all reefs. However, we found that the coral reef fish composition remained largely stable through time and space whereas compensatory changes in biomass among species were responsible for most of the temporal fluctuations, as outlined by the overall high contribution of the replacement component to total beta diversity. This suggests that, despite the severity of the two disturbances, fish communities exhibited high resistance and the ability to reorganize their compositions to maintain the same level of total community biomass as before the disturbances. We further investigated the spatial congruence of this pattern and showed that temporal dynamics involved different species across sites; yet, herbivores controlling the proliferation of algae that compete with coral communities were consistently favored. These results suggest that compensatory changes in biomass among species and spatial heterogeneity in species responses can provide further insurance against natural disturbances in coral reef ecosystems by promoting high levels of key species (herbivores). They can also allow the ecosystem to recover more quickly.     

Timescales of population rarity and commonness in random environments

This is a mathematical study of the interactions between non-linear feedback (density dependence) and uncorrelated random noise in the dynamics of unstructured populations. The stochastic non-linear dynamics are generally complex, even when the deterministic skeleton possesses a stable equilibrium. There are three critical factors of the stochastic non-linear dynamics; whether the intrinsic population growth rate (lambda) is smaller than, equal to, or greater than 1; the pattern of density dependence at very low and very high densities; and whether the noise distribution has exponential moments or not. If lambda < 1, the population process is generally transient with escape towards extinction. When lambda > or = 1, our quantitative analysis of stochastic non-linear dynamics focuses on characterizing the time spent by the population at very low density (rarity), or at high abundance (commonness), or in extreme states (rarity or commonness). When lambda >1 and density dependence is strong at high density, the population process is recurrent: any range of density is reached (almost surely) in finite time. The law of time to escape from extremes has a heavy, polynomial tail that we compute precisely, which contrasts with the thin tail of the laws of rarity and commonness. Thus, even when lambda is close to one, the population will persistently experience wide fluctuations between states of rarity and commonness. When lambda = 1 and density dependence is weak at low density, rarity follows a universal power law with exponent -3/2. We provide some mathematical support for the numerical conjecture [Ferriere, R., Cazelles, B., 1999. Universal power laws govern intermittent rarity in communities of interacting species. Ecology 80, 1505-1521.] that the -3/2 power law generally approximates the law of rarity of 'weakly invading' species with lambda values close to one. Some preliminary results for the dynamics of multispecific systems are presented.     

A general framework for neutral models of community dynamics

Neutral models of community dynamics are a powerful tool for ecological research, but their applications are currently limited to unrealistically simple types of dynamics and ignore much of the complexity that characterize natural ecosystems. Here, we present a new analytical framework for neutral models that unifies existing models of neutral communities and extends the applicability of existing models to a much wider spectrum of ecological phenomena. The new framework extends the concept of neutrality to fitness equivalence and in spite of its simplicity explains a wide spectrum of empirical patterns of species diversity including positive, negative and unimodal productivity–diversity relationships; gradual and highly delayed declines in species diversity with habitat loss; and positive and negative responses of species diversity to habitat heterogeneity. Surprisingly, the abundance distribution in all of these cases is given by the dispersal limited multinomial (DLM), the abundance distribution in Hubbell's zero-sum model, showing DLM's robustness and demonstrating that it cannot be used to infer the underlying community dynamics. These results support the hypothesis that ecological communities are regulated by a limited set of fundamental mechanisms much simpler than could be expected from their immense complexity.     

Species abundance distributions as a proxy for the niche-neutrality continuum

Aims Species abundance distributions (SADs) are often used to verify mechanistic theories underlying community assembly. However, it is now accepted that SADs alone are not sufficient to reveal biological mechanisms. Recent attention focuses on the relative importance of stochastic dispersal processes versus deterministic processes such as interspecific competition and environmental filtering. Here, we combine a study of the commonness and rarity of species (i.e. the SAD) with mechanistic processes underlying community composition. By comparing the occurrence frequencies of each and every species with its abundance, we quantify the relative contributions of common and rare species to the maintenance of community structure. Essentially, we relate the continuum between commonness and rarity with that of niches and neutrality.Methods An individual-based, spatially explicit model was used to simulate local communities in niche spaces with the same parameters. We generated sets of assemblages from which species were eliminated in opposing sequences: from common to rare and from rare to common, and investigated the relationship between the abundance and frequency of species. We tested the predictions of our model with empirical data from a field experiment in the environmentally homogeneous alpine meadows of the Qinghai–Tibetan plateau.Important findings Our simulations support the widespread notion that common species maintain community structure, while rare species maintain species diversity, in both local and regional communities. Our results, both from theoretical simulations and from empirical observations, revealed positive correlations between the abundance of a particular species and its occurrence frequency. SAD curves describe a continuum between commonness and rarity. Removing species from the 'rare' end of this continuum has little effect on the similarity of communities, but removing species from the 'common' end of the continuum causes significant increases in beta diversity, or species turnover, between communities. In local communities distributed in a homogenous habitat, species located at the 'common' end of the continuum should be selected by environmental filtering, with niche space partitioning governed by interspecific competition. Conversely, species located at the 'rare' end of the continuum are most likely subject to stochastic dispersal processes. Species situated at intermediate locations on this continuum are therefore determined by niche and neutral processes acting together. Our results suggest that, in homogeneous habitats, SAD curves describing the common-rare continuum may also be used to describe the continuum between niches and neutrality.     

Evaluating functional diversity conservation for freshwater fishes resulting from terrestrial protected areas

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The impact of neutrality, niche differentiation and species input on diversity and abundance distributions

We present a spatially-explicit generalization of Hubbell's model of community dynamics in which the assumption of neutrality is relaxed by incorporating dispersal limitation and habitat preference. In simulations, diversity and species abundances were governed by the rate at which new species were introduced (usually called 'speciation') and nearly unaffected by dispersal limitation and habitat preference. Of course, in the absence of species input, diversity is maintained solely by niche differences. We conclude that the success of the neutral model in predicting the abundance distribution has nothing to do with neutrality, but rather with the species-introduction process: when new species enter a community regularly as singletons, the typical J-shaped abundance distribution, with a long tail of rare species, is always observed, whether species differ in habitat preferences or not. We suggest that many communities are indeed driven by the introduction process, accounting for high diversity and rarity, and that species differences may be largely irrelevant for either.     

Stochastic species turnover and stable coexistence in a species-rich, fire-prone plant community

Understanding the mechanisms that maintain diversity is important for managing ecosystems for species persistence. Here we used a long-term data set to understand mechanisms of coexistence at the local and regional scales in the Cape Floristic Region, a global hotspot of plant diversity. We used a dataset comprising 81 monitoring sites, sampled in 1966 and again in 1996, and containing 422 species for which growth form, regeneration mode, dispersal distance and abundances at both the local (site) and meta-community scales are known. We found that species presence and abundance were stable at the meta-community scale over the 30 year period but highly unstable at the local scale, and were not influenced by species' biological attributes. Moreover, rare species were no more likely to go extinct at the local scale than common species, and that alpha diversity in local communities was strongly influenced by habitat. We conclude that stochastic environmental fluctuations associated with recurrent fire buffer populations from extinction, thereby ensuring stable coexistence at the meta-community scale by creating a "neutral-like" pattern maintained by niche-differentiation.     

The contribution of rare species to community phylogenetic diversity across a global network of forest plots

Niche differentiation has been proposed as an explanation for rarity in species assemblages. To test this hypothesis requires quantifying the ecological similarity of species. This similarity can potentially be estimated by using phylogenetic relatedness. In this study, we predicted that if niche differentiation does explain the co-occurrence of rare and common species, then rare species should contribute greatly to the overall community phylogenetic diversity (PD), abundance will have phylogenetic signal, and common and rare species will be phylogenetically dissimilar. We tested these predictions by developing a novel method that integrates species rank abundance distributions with phylogenetic trees and trend analyses, to examine the relative contribution of individual species to the overall community PD. We then supplement this approach with analyses of phylogenetic signal in abundances and measures of phylogenetic similarity within and between rare and common species groups. We applied this analytical approach to 15 long-term temperate and tropical forest dynamics plots from around the world. We show that the niche differentiation hypothesis is supported in six of the nine gap-dominated forests but is rejected in the six disturbance-dominated and three gap-dominated forests. We also show that the three metrics utilized in this study each provide unique but corroborating information regarding the phylogenetic distribution of rarity in communities.     

Synchrony in small mammal community dynamics across a forested landscape

Long‐term studies at local scales indicate that fluctuations in abundance among trophically similar species are often temporally synchronized. Complementary studies on synchrony across larger spatial extents are less common, as are studies that investigate the subsequent impacts on community dynamics across the landscape. We investigate the impact of species population fluctuations on concordance in community dynamics for the small mammal fauna of the White Mountain National Forest, USA. Hierarchical open population models, which account for imperfect detection, were used to model abundance of the most common species at 108 sites over a three year period. Most species displayed individualistic responses of abundance to forest type and physiographic characteristics. However, among species, we found marked synchrony in population fluctuations across years, regardless of landscape affinities or trophic level. Across the region, this population synchrony led to high within‐year concordance of community composition and aggregate properties (e.g. richness and diversity) independent of forest type and low among‐year similarity in communities, even for years with similar species richness. Results suggest that extrinsic factors primarily drive abundance fluctuations and subsequently community dynamics, although local community assembly may be modified by species dispersal abilities and biotic interactions. Concordant community dynamics across space and over time may impact the stability of regional food webs and ecosystem functions.     

Herbivory and drought interact to enhance spatial patterning and diversity in a savanna understory

The combination of abiotic stress and consumer stress can have complex impacts on plant community structure. Effective conservation and management of semi-arid ecosystems requires an understanding of how different stresses interact to structure plant communities. We explored the separate and combined impacts of episodic drought, livestock grazing, and wild ungulate herbivory on species co-occurrence and diversity patterns in a relatively productive, semi-arid Acacia savanna. Specifically, we analyzed 9 years of biannual plant community data from the Kenya long-term exclosure experiment, a broad-scale manipulative experiment that has excluded different combinations of large mammalian herbivores from 18 4-ha plots since 1995. During droughts, we observed low species diversity and random species co-occurrence patterns. However, when rain followed a major drought, areas exposed to moderate cattle grazing displayed high species diversity and evidence of significant species aggregation. These patterns were not apparent in the absence of cattle, even if other large herbivores were present. To explore possible mechanisms, we examined patterns separately for common and rare species. We found that aggregation patterns were likely driven by rare species responding similarly to the availability of open micro-sites. Our results indicate that in a productive, fire-suppressed savanna, the combination of periodic drought and moderate cattle grazing can enhance plant biodiversity and fine-scale spatial heterogeneity by opening up space for species that are otherwise rare or cryptic. Our findings also emphasize that domestic herbivores can have significantly stronger impacts on plant community dynamics than wild herbivores, even in an ecosystem with a long history of grazing.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

The functional importance of rare and dominant species in a Neotropical forest bird community

Bird communities in Tropical forests have high rate of rare species, but only recently some studies suggested their disproportional role for the overall functional diversity. We investigated data from bird communities monitored annually using point counts over ten years in Iguassu National Park, that is a large and protected area in the southern portion of the Atlantic Forest in Brazil. We aimed (1) to determine the rare and dominant species in the community based on their abundance over time; (2) to evaluate the impact of each class of rarity on the overall functional volume of the community; (3) to investigate the pattern of occupancy of the trait space filled by each dominance class, and (4) to assess the disparity in functional trait composition between classes of rarity. We defined dominant, intermediate, and rare species in communities using cluster analysis and data of relative abundance of species in five sections of 1-km in the forest interior. The number of clusters was defined in accordance with the silhouette criterion measures of cohesion and separation that range from −1 to 1, with values > 0.5 indicating high-quality clusters. Among total 138 bird species recorded, 107 were rare, 26 intermediates, and 5 dominants. Our data corroborate a functional disproportional importance of rare bird species in the community considering the functional space (FRic), but functional dispersion (FDis) was not significantly different between the rare and intermediate species. In fact, there is a large overlap of the functional volume occupied by rare in relation to dominant/intermediate species. The niche space occupied by rare species covers most of the space occupied by intermediate and dominant species. The low representativeness of functional turnover indicates that few functions are unique to the classes of higher dominance. Our data suggest the importance of rare bird species to the overall functional diversity but also highlights the potential use of dominants/intermediates species as indicators to select important forests areas for conservation, as certain forest fragments. Through these most abundant species it would be possible to assess which functions are heaviest in abundance, becoming core functions, in the context of each independent forest fragment.