Historical contingency in species interactions: towards niche‐based predictions

The way species affect one another in ecological communities often depends on the order of species arrival. The magnitude of such historical contingency, known as priority effects, varies across species and environments, but this variation has proven difficult to predict, presenting a major challenge in understanding species interactions and consequences for community structure and function. Here, we argue that improved predictions can be achieved by decomposing species' niches into three components: overlap, impact and requirement. Based on classic theories of community assembly, three hypotheses that emphasise related, but distinct influences of the niche components are proposed: priority effects are stronger among species with higher resource use overlap; species that impact the environment to a greater extent exert stronger priority effects; and species whose growth rate is more sensitive to changes in the environment experience stronger priority effects. Using nectar‐inhabiting microorganisms as a model system, we present evidence that these hypotheses complement the conventional hypothesis that focuses on the role of environmental harshness, and show that niches can be twice as predictive when separated into components. Taken together, our hypotheses provide a basis for developing a general framework within which the magnitude of historical contingency in species interactions can be predicted.     

Light interception in species with different functional groups coexisting in moorland plant communities

Competition for light is one of the most essential mechanisms affecting species composition. It has been suggested that similar light acquisition efficiency (Φ_mass, absorbed photon flux per unit aboveground mass) may contribute to species coexistence in multi-species communities. On the other hand, it is known that traits related with light acquisition vary among functional groups. We studied whether Φ_mass was similar among species with different functional groups coexisting in moorland communities. We conducted stratified clipping in midsummer when the stand biomass reached a maximum. Light partitioning among species was estimated using a model accounting for both direct and diffuse light. Evergreen species were found to have a significantly lower Φ_mass than deciduous species, which resulted from their lower absorbed photon flux per unit leaf area and lower specific leaf area. Shrubs had a smaller leaf mass fraction, but their Φ_mass was not lower than that of herbs because they had a higher leaf position due to the presence of wintering stems. Species with vertical leaves had a higher Φ_mass than those with horizontal leaves despite vertical leaves being a decided disadvantage in terms of light absorption. This higher Φ_mass was achieved by a greater leaf height in species with vertical leaves. Our results clearly demonstrate that light acquisition efficiency was different among the functional groups. However, the trend observed is not necessarily the same as that expected based on prior knowledge, suggesting that disadvantages in some traits for light acquisition efficiency are partly compensated for by other traits.     

Hypervolume concepts in niche‐ and trait‐based ecology

Hutchinson's n‐dimensional hypervolume concept for the interpretation of niches as geometric shapes has provided a foundation for research across different fields of ecology and evolution. There is now an expanding set of applications for hypervolume concepts, as well as a growing set of statistical methods available to operationalize this concept with data. The concept has been applied to environmental, resource, functional trait, and morphometric axes and to different scales, i.e. from individuals, species, to communities and clades. Further, these shapes have been variously interpreted as niches, ecological or evolutionary strategy spaces, or proxies for community structure. This paper highlights these applications’ shared mathematical framework, surveys uses of the hypervolume concept across fields, discusses key limitations and assumptions of hypervolume concepts in general, provides a critical guide to available statistical estimation methods, and delineates the situations where hypervolume concepts can be useful.     

Fire‐mediated niche‐separation between two sympatric small mammal species

Fire is a key ecological process influencing the population dynamics of small mammals. Whilst shifting competitive advantage amongst small mammal species following a single fire event is well‐documented, there has been little investigation of the potential influence of fire frequency on small mammal interspecific interactions. In this study, we investigated the effect of fire frequency on the abundance of two small dasyurid mammals, Antechinus stuartii and A. flavipes, which occur sympatrically in some parts of their range. The two antechinus species are known to have different habitat preferences, so it is possible that fire regimes may promote their coexistence in areas of sympatry by altering vegetation structure. To investigate this possibility, we estimated the abundance of both species using replicate sites which differed in the number of times burnt (1–4) during the last four decades, but with identical time‐since‐fire. Proportionally, we captured greater numbers of A. stuartii in less frequently burnt sites and greater numbers of A. flavipes in more‐frequently burnt sites. Hence, fire may mediate niche‐separation between these two species. To clarify further this pattern of response to fire frequency, we investigated which structural habitat variables differed between fire frequencies, and compared antechinus abundances with structural vegetation characteristics. We found a trend for lower ground cover density under higher fire frequencies. This offers one potential explanation of the patterns of abundance that we observed. Our study provided insights into the complexities of small mammal responses to fire, and strongly suggests that fire could mediate competitive interactions between species.     

Diversity partitioning of moorland plant communities across hierarchical spatial scales

Understanding of the scaling of diversity is critical to enhance conservation strategies for subalpine moorland ecosystems vulnerable to future environmental changes. However, a paucity of quantitative data strongly limits such attempts. In this study, we used an additive diversity partitioning framework and quantified diversity patterns of moorland plant communities across hierarchical spatial scales, within- and between-sample transects, and between sites (corresponding to α and two levels of β diversity). Moorland sites markedly differed in size (range 1,000–160,000 m²) and were isolated from each other to varying extents within an inhospitable matrix (i.e., forests). We found that β diversity components were consistently higher, whereas the local α diversity component was consistently lower than expected by chance. We observed substantial contribution at the between-site scale to total species richness. By focusing on diversity patterns of moorland plant communities across multiple hierarchical spatial scales, we could thus identify the scale at which regional diversity is maximized. Our results suggest that protection of as many moorland sites as possible, to ensure beta diversity between sites, will effectively conserve total diversity. The use of additive diversity partitioning is a major step forward in providing strategies for the biological conservation of subalpine moorland ecosystems vulnerable to future environmental changes.     

Functional trade‐offs increase species diversity in experimental plant communities

Functional trade‐offs have long been recognised as important mechanisms of species coexistence, but direct experimental evidence for such mechanisms is extremely rare. Here, we test the effect of one classical trade‐off – a negative correlation between seed size and seed number – by establishing microcosm plant communities with positive, negative and no correlation between seed size and seed number and analysing the effect of the seed size/number correlation on species richness. Consistent with theory, a negative correlation between seed size and seed number led to a higher number of species in the communities and a corresponding wider range of seed size (a measure of functional richness) by promoting coexistence of large‐ and small‐seeded species. Our study provides the first direct evidence that a seed size/number trade‐off may contribute to species coexistence, and at a wider context, demonstrates the potential role of functional trade‐offs in maintaining species diversity.     

Unprotecting the rare species: a niche‐based gap analysis for odonates in a core Cerrado area

Aim We evaluated Odonata distribution data and predicted the compositional resemblance based on niche‐based species distribution models to analyse the following questions: (1) How is estimated species richness distributed, and how can it be preserved under the actual network of conservation units (a gap analysis approach)? (2) How is the estimated odonate beta diversity distributed, and is there a better distribution of conservation units (a priority setting approach)? (3) Is the probability of being under protection a function of the potential species range size? and (4) Will the current conservation network proposals protect odonate taxa? Location Central Brazil in a core Cerrado area. Methods We generated odonate species distribution predictions based on MaxEnt and maps derived from estimated species richness, beta diversity and gap analysis for all species predicted to occur in the study area. Then, we compared these maps with current conservation units, land‐use patterns and proposals for the establishment of conservation units. Results Raw odonate species records provided limited utility for setting conservation priorities without the use of niche‐based models. However, area under the receiver operating curve (AUC) values were characterized by substantial variation that was related to the number of records. No current conservation units overlapped the areas with higher predicted richness and beta diversity, and in general, conservation units were not preserving restricted/rare species. There was a direct linear correlation between species range size and the proportion of its range protected in the current network of conservation units. Finally, we identified three areas with high odonate beta diversity where conservationist actions should be implemented. Main conclusions Current conservation units and future suggested areas do not overlap regions with high conservation values for odonates. Conservation units protect species at random, and the level of protection has a direct relationship with species range size; thus, wide‐range species are expected to be more protected than restricted or threatened species.     

Can niche‐based distribution models outperform spatial interpolation?

Aim Distribution modelling relates sparse data on species occurrence or abundance to environmental information to predict the population of a species at any point in space. Recently, the importance of spatial autocorrelation in distributions has been recognized. Spatial autocorrelation can be categorized as exogenous (stemming from autocorrelation in the underlying variables) or endogenous (stemming from activities of the organism itself, such as dispersal). Typically, one asks whether spatial models explain additional variability (endogenous) in comparison to a fully specified habitat model. We turned this question around and asked: can habitat models explain additional variation when spatial structure is accounted for in a fully specified spatially explicit model? The aim was to find out to what degree habitat models may be inadvertently capturing spatial structure rather than true explanatory mechanisms. Location We used data from 190 species of the North American Breeding Bird Survey covering the conterminous United States and southern Canada. Methods We built 13 different models on 190 bird species using regression trees. Our habitat‐based models used climate and landcover variables as independent variables. We also used random variables and simulated ranges to validate our results. The two spatially explicit models included only geographical coordinates or a contagion term as independent variables. As another angle on the question of mechanism vs. spatial structure we pitted a model using related bird species as predictors against a model using randomly selected bird species. Results The spatially explicit models outperformed the traditional habitat models and the random predictor species outperformed the related predictor species. In addition, environmental variables produced a substantial R² in predicting artificial ranges. Main conclusions We conclude that many explanatory variables with suitable spatial structure can work well in species distribution models. The predictive power of environmental variables is not necessarily mechanistic, and spatial interpolation can outperform environmental explanatory variables.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

Prediction of species composition of plant communities in a rural landscape based on species traits

The species composition of a community is a subset of the regional species pool, and predicting the species composition of a community from ecological traits of organisms is an important objective in ecology. If such a prediction can be made feasible, we could assess the risk of invasion of locally new species (alien species and genetically modified species) into natural communities. We developed and tested statistical models to predict a community’s species composition from ecological traits of the species pool. Various types of communities (forest, meadow, and weed communities) exist in a small area of traditional rural landscape in Japan, and have been maintained by human activities. These communities and the tracheophytes species pool in the 1-km² research area were considered. We used logistic regression and decision-tree analysis to construct predictive models of community species composition based on plant traits, using the presence or absence of species in a community as the dependent variable and ecological traits as independent variables. Plant traits were grouped by cluster analysis, and the average in each trait group was used for model building to avoid multiple collinearity. Statistical prediction models were significant in all communities. About 60–75% of species composition could be predicted from the measured plant traits in forest communities, but 33–56% in the meadow and weed communities. Our results showed the possibility of predicting the species composition of plant communities from the ecological traits of the plant species together with the information on local species pool.     

From species distributions to meta‐communities

The extent that biotic interactions and dispersal influence species ranges and diversity patterns across scales remains an open question. Answering this question requires framing an analysis on the frontier between species distribution modelling (SDM), which ignores biotic interactions and dispersal limitation, and community ecology, which provides specific predictions on community and meta‐community structure and resulting diversity patterns such as species richness and functional diversity. Using both empirical and simulated datasets, we tested whether predicted occurrences from fine‐resolution SDMs provide good estimates of community structure and diversity patterns at resolutions ranging from a resolution typical of studies within reserves (250 m) to that typical of a regional biodiversity study (5 km). For both datasets, we show that the imprint of biotic interactions and dispersal limitation quickly vanishes when spatial resolution is reduced, which demonstrates the value of SDMs for tracking the imprint of community assembly processes across scales.     

Sugar landscapes and pollinator‐mediated interactions in plant communities

Pollinator‐mediated interactions between plants can play an important role for the dynamics of plant communities. Pollination services depend on the abundance and the foraging behaviour of pollinators, which in turn respond to the availability and distribution of floral resources (notably nectar sugar). However, it is still insufficiently understood how the ‘sugar landscapes’ provided by flowering plant communities shape pollinator‐mediated interactions between multiple plant species and across different spatial scales. A better understanding of pollinator‐mediated interactions requires an integrative approach that quantifies different aspects of sugar landscapes and investigates their relative importance for pollinator behaviour and plant reproductive success. In this study, we quantified such sugar landscapes from individual‐based maps of Protea shrub communities in the Cape Floristic Region, South Africa. The 27 study sites of 4 ha each jointly comprise 127 993 individuals of 19 species. We analysed how rates of visitation by key bird pollinators and the seed set of plants respond to different aspects of sugar landscapes: the distribution of nectar sugar amounts, as well as their quality, taxonomic purity and phenology. We found that pollinator visitation rates strongly depended on phenological variation of site‐scale sugar amounts. The seed set of focal plants increased with nectar sugar amounts of conspecific neighbours and with site‐scale sugar amounts. Seed set increased particularly strongly if site‐scale sugar amounts were provided by plants that offer less sugar per inflorescence. These combined effects of the amount, quality, purity and phenological variation of nectar sugar show that nectar sugar is a common interaction currency that determines how multiple plant species interact via shared pollinators. The responses of pollinator‐mediated interactions to different aspects of this interaction currency alter conditions for species coexistence in Protea communities and may cause community‐level Allee effects that promote extinction cascades.     

Resource availability determines the importance of niche‐based versus stochastic community assembly in grasslands

Niche‐based selection and stochastic processes can operate simultaneously to generate spatial and temporal variation in species composition. Yet, the conditions under which ecological dynamics are dominated by niche‐based versus stochastic processes are poorly understood. Using a field experiment in early‐successional temperate grassland and null models of beta diversity, this study investigates the effects of soil nutrient supply on the relative importance of niche‐based selection versus stochastic dynamics for variation in species composition among sites. Nutrient availability was manipulated experimentally, individual seed mixtures with 25 species were sown in each experimental plot, and then stochastic and deterministic niche‐based assembly processes were allowed to happen. We found that compositional variation among grassland plots with low nutrient supply was driven by stochastic immigration and extinctions. In contrast, nutrient enrichment reduced the importance of stochasticity and imposed a deterministic environmental filter that homogenized communities through the selection of few species with greater competitive ability for light. This demonstrates that soil nutrient availability is a critical environmental feature that dictates the degree to which terrestrial plant communities are controlled by niche‐based selection versus stochastic assembly processes. Our study shows further that alternative states of eutrophic grasslands emerge from initial stochastic variation in the composition of a particular functional group of species that can become dominant at high nutrient supply. We discuss potential mechanisms underlying the shift from stochastic to niche‐driven dynamics along soil nutrient gradients.     

Spatial patterns and species performances in experimental plant communities

The roles of co-occurring herbivores that modify habitat structure and ecosystem processes have seldom been examined in manipulative experiments or explored in early successional communities. In a created marsh in southern California (USA), we tested the individual and combined effects of two epibenthic invertebrates on nutrient and biomass pools, community structure, and physical habitat features. We manipulated snail (Cerithidea californica) and crab (Pachygrapsus crassipes) presence in field enclosures planted with pickleweed (Salicornia virginica) at elevations matching the plant’s lower extent in an adjacent natural marsh. In the 4-month experiment, C. californica altered habitat structure by reducing sediment surface heterogeneity and shear strength (a measure of sediment stability) markedly throughout the enclosures. Both invertebrates had strong negative effects on a group of correlated sediment physicochemical characteristics, including nitrogen and organic matter concentrations and soil moisture. In addition, both invertebrates greatly reduced benthic chlorophyll a, a proxy for biomass of microphytobenthos. Compared to controls, macroalgal cover was up to sixfold lower with crabs present, while snails increased cover at low elevations of enclosures. Unexpectedly, macroalgal cover was eliminated with both species present, perhaps through P. crassipes consumption of larger thalli and C. californica reduction in cover of recruits. Neither species influenced the S. virginica canopy (quantified with an index of branch length and number); however, at the lower elevation of enclosures, the two species together negatively impacted the plant canopy. The two invertebrates’ modifications to our experimental marshes led to distinct suites of biotic and physicochemical features depending on their presence or co-occurrence, with the latter producing several unexpected results. We propose that the roles and interactions of habitat-modifying fauna deserve further attention, particularly in the context of efforts to conserve and restore the processes found in natural systems.     

The phylogenetic properties of native‐ and exotic‐dominated plant communities

Phylogenetic properties of communities (phylogenetic diversity and phylogenetic structure) allow for the characterisation of phylogenetic patterns and provide the information necessary to infer mechanisms of species assembly. Because humans have introduced exotic species and modified the physical conditions of landscapes, the phylogenetic properties of communities should change according to the proportion of natives to exotics hosted by sites and to the strength of the conditions that act as habitat filters in human‐disturbed habitats. To assess the effects of the introduction of exotic plant species, we characterized the phylogenetic properties of 67 plant communities with different degrees of exotic species dominance in a region of central Chile with a Mediterranean climate. Five indices were used to estimate the phylogenetic properties. The Faith index (FPD), the mean pairwise distance (MPD) and the mean nearest neighbour distance (MNND) were used to estimate phylogenetic diversity, and the nearest relative index (NRI) and the nearest taxon index (NTI) were used as estimators of the phylogenetic structure (the phylogenetic distribution of taxa in a community) of species assemblages. We observed greater phylogenetic diversity of natives versus exotic plants despite the fact that natives accounted for a fewer number of taxa among the studied communities. Second, assemblages exhibited a phylogenetically clustered structure, which is attributable to an over‐representation of some families of exotic flora (Asteraceae, Brassicaceae, Fabaceae, Papaveraceae, Poaceae) and suggests habitat filtering processes that could have acted by selecting species with traits that permit adaptation to the harsh conditions of human‐disturbed sites.     

陆地植物群落物种多样性研究进展

生物多样性是当前生态学研究的热点之一,物种多样性层次是最直接、最易观察和最适合研究生物多样性的层次。总结了与群落动态、生境因子、取样尺度及生态系统相关的陆地植物物种多样性研究。同时,根据目前的趋势提出了多样性动态研究的发展动向。     

Nestedness in riverine mussel communities: patterns across sites and fish hosts

The pattern of nestedness, where species present in depauperate locations are subsets of species present in locations with higher species diversity, is often found in ecological communities. Mussel communities examined in four rivers in the upper Tennessee River basin appeared significantly nested. Mussel species distributions were mostly unrelated to differences in immigration and only weakly related to downstream direction, giving some indication of structuring by differences in extinction. Mussel species distributions were not related to the number of fish species used as hosts for mussel larvae. Mussel species were more likely to overlap on common fish hosts; however, the host‐use matrix was not nested – groups of mussel species used different sets of host fish species in a pattern that appeared phylogenetically related. Sites with high fish host abundance may support high mussel diversity by promoting the survival of mussel species that are less able to attract and infect hosts. Thus, nestedness in freshwater mussel communities may be driven by the array of host fish resources, combined with differences in species’ abilities to use fish hosts. An understanding of the nested pattern in this region can aid conservation of this imperiled fauna.     

Are bryophyte communities different from higher‐plant communities? Abundance relations

The considerable differences in biology between bryophytes and higher plants have led to speculation that their community structure might be different. Ten bryophyte communities were sampled for species biomass composition, and for comparison ten higher-plant communities that were similar in physiognomy and in total community biomass. The rather insecure theory in the bryophyte literature was distilled into eight quantifiable predictions, which were tested. For seven, there was no sign of the predicted differences: i.e. no indication of the predicted low within-community heterogeneity, higher species richness, more variable species richness, lower rank consistency, a poor fit for the geometric model of RAD (relative abundance distribution), better fit for the broken-stick and general-lognormal RAD models with general-lognormal parameter γ deviating further from 1.0, or of a good fit for the Zipf-Mandelbrot RAD model. However, evenness was, on average, significantly (p=0.005) less in the bryophyte communities, using any of four evenness indices. Two possible features of bryophytes are suggested that might cause this: (a) a smaller module (i.e. shoot, leaf) size, allowing species to be present with a lower threshold biomass, and (b) less efficient competitive exclusion among bryophytes because of weaker competition and a predominance of mutualism, as suggested in the literature. However, the striking conclusion from the results is that in spite of all the biological differences between the two groups of organisms, their community organisation is remarkably similar.