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.     

Nutrient availability and grazing influence the strength of priority effects during freshwater bacterial community coalescence

When bacterial communities mix, immigration history can fundamentally affect the community composition as a result of priority effects. Priority effects arise when an early immigrant exhausts resources and/or alters habitat conditions, thereby influencing the establishment success of the late arriver. The strength of priority effects is context-dependent and expected to be stronger if environmental conditions favour the growth of the first arriver. In this study, we conducted a two-factorial experiment testing the importance of nutrient availability and grazing on the strength of priority effects in complex aquatic bacterial communities. We did so by mixing two dissimilar communities, simultaneously, and with a 38 h time-delay. Priority effects were measured as the invasion resistance of the first community to the invading second community. We found stronger priority effects in treatments with high nutrient availability and absence of grazing, but in general, the arrival timing was less important than the selection by nutrients and grazing. At the population level, the results were complex, but priority effects may have been driven by bacteria belonging to for example, the genera Rhodoferax and Herbaspirillum. Our study highlights the importance of arrival timing in complex bacterial communities, especially if environmental conditions favour rapid community growth.     

Resource composition mediates the effects of intraspecific variability in nutrient recycling on ecosystem processes

Despite the growing evidence for individual variation in trophic niche within populations, its potential indirect effects on ecosystem processes remains poorly understood. In particular, few studies have investigated how intraspecific trophic variability can modulate the effects of consumers on ecosystems through potential changes in nutrient excretion rates. Here, we first quantified the level of intraspecific trophic variability in 11 wild populations of the omnivorous fish Lepomis gibbosus. Outputs from stomach content and stable isotope analyses revealed that the degree of trophic specialization and trophic positions were highly variable between and within these wild populations. There was intrapopulation variation in trophic position of more than one trophic level, suggesting that individuals consumed a range of plant and animal resources. We then experimentally manipulated intraspecific trophic variability to assess how it can modulate consumer‐mediated nutrient effects on relevant processes of ecosystem functioning. Specifically, three food sources varying in nutrient quality (e.g. plant material, macro‐invertebrate and fish meat) were used individually or in combination to simulate seven diet treatments. Results indicated that intraspecific variability in growth and nitrogen excretion rates were more related to the composition of the diet rather than the degree of specialization, and increased with the trophic position of the diet consumed. We subsequently used microcosms and showed that critical ecosystem functions, such as primary production and community respiration, were affected by the variability in excretory products, and this effect was biomass‐dependent. These results highlight the importance of considering variation within species to better assess the effects of individuals on ecosystems and, more specifically, the effects of consumer‐mediated nutrient recycling because the body size and the trophic ecology of individuals are affected by a large spectrum of natural and human‐induced environmental changes.     

Resource availability dominates and alters the relationship between species diversity and ecosystem productivity in experimental plant communities

Experimental evidence that plant species diversity has positive effects on biomass production appears to conflict with correlations of species diversity and standing biomass in natural communities. This may be due to the confounding effects of a third variable, resource availability, which has strong control over both diversity and productivity in natural systems and may conceal any positive effects of diversity on productivity. To test this hypothesis, I independently manipulated resource availability (soil fertility) and sown species diversity in a field experiment and measured their individual and interactive effects on productivity. Although fertility was a far stronger predictor of productivity than diversity, the effect of diversity on productivity significantly increased with fertility. Relative yield analyses indicated that plant mixtures of high fertility treatments significantly "overyielded," or were more productive than expected based on monoculture yields of component species. In contrast, plant mixtures of low fertility treatments had significantly lower-than-expected yields. The effect of diversity on productivity was also driven by sampling effects, where more species-rich mixtures were more likely to include particularly productive species. Unexpectedly, the strength of sampling effects was largely insensitive to fertility, although the particular species most responsible for sampling effects did change with fertility. These results suggest that positive effects of species diversity on ecosystem productivity in natural systems are likely to be masked by variation in environmental factors among habitats.     

Interactions after death: plant litter controls priority effects in a successional plant community

We performed a field experiment to test whether the presence of litter produced by the dominant species in the first successional year affects the plant community structure in the following year. We removed the litter of Setaria faberii (the first-year dominant) in midfall, early spring, mid-spring, or late spring. Both the fall and early spring removal increased the biomass of Erigeron annuus, which became dominant, and reduced the biomass of S. faberii. In the fall-removal treatment more plants of E. annuus flowered, while early spring removal increased the biomass of rosettes (non-flowering individuals) at the end of the growing season. In the other treatments and in the control S. faberii retained dominance, but its biomass was the highest in mid-spring removal plots. The removal of litter of S. faberii in the fall and in early spring allowed E. annuus to pre-empt the site and dominate the community. When litter was not removed, it strongly hindered the growth of E. annuus, favoring S. faberii. These results highlight the importance of litter as a historical factor linking interactions across successive generations, and controlling the community structure.     

Effects of resource availability on plant recruitment at the community level in a Mediterranean mountain ecosystem

Coexisting plant species usually differ in resource requirements, which may also vary within species at successive demographic stages. Such differences become extremely important during the early life stages, since these are the most critical phases in woody-species recruitment, they depend heavily on resources, and they may determine future community composition. Under a global-change scenario, where climatic conditions, nutrient availability, and habitat characteristics are expected to be altered, it is difficult to predict the way in which plant recruitment will be affected. To understand the impact of different global-change drivers on community recruitment, we sowed a set of species representative of the different successional groups of a complete Mediterranean woody community under field conditions, and studied their emergence, growth, and survival along the main resource gradients of light, water, and nutrients. The light and nutrient gradients followed the natural range of conditions in the study area, but water availability was manipulated to simulate three contrasting climatic scenarios: wetter, drier, and current conditions. Structural equation modelling was used to provide a comprehensive analysis of the factors and relations governing plant recruitment. Overall, seedling emergence was determined directly by light; growth was determined by light and summer soil moisture; and survival was determined by summer soil moisture. Light was the main factor indirectly affecting the demographic stages of all species. However, the magnitude of the direct and indirect relationships varied among species. Particularly, species differed in their response to the expected drier climatic conditions, some (e.g. Pinus sylvestris, Acer opalus) being much more vulnerable than others (e.g. Cytisus scoparius, Salvia lavandulifolia). These differential responses could translate as major shifts in the structure of the overall plant community. Our results support the idea that the analysis of complex relations among essential resources is critical for accurate forecasts of the impact of climate change on community dynamics.     

Resource dependence in a new ecosystem: A host plant and its colonizing community

The introduced black locust (Robinia pseudoacacia) has become an invasive plant species in Europe. The introduction of alien plants such as the black locust may modify ecosystem composition and functioning. In response to the presence of a potential host plant, herbivores can adapt and shift to the consumption of the new host plant. In Eastern-Central Europe, the seed predator Bruchophagus robiniae (Hymenoptera: Eurytomidae) is an important consumer of black locust seeds which presumably shifted from its formerly host species to black locust. We tested the influence of host plant abundance on a seed predator – parasitoid community. We found that the seed predator B. robiniae was present in higher numbers in woodlots than in small patches of black locust. The density of the specialist parasitoid Mesopolobus sp. was lower in woodlots than in small patches, while the generalist parasitoid Eupelmus urozonos was evenly distributed between woodlots and small patches of black locust. We found that parasitoid species are influenced by the patch size of host plants, thus characteristics of introduced host plants can also manifest in higher trophic levels.     

Climate change drives a shift in peatland ecosystem plant community: Implications for ecosystem function and stability

The composition of a peatland plant community has considerable effect on a range of ecosystem functions. Peatland plant community structure is predicted to change under future climate change, making the quantification of the direction and magnitude of this change a research priority. We subjected intact, replicated vegetated poor fen peat monoliths to elevated temperatures, increased atmospheric carbon dioxide (CO₂), and two water table levels in a factorial design to determine the individual and synergistic effects of climate change factors on the poor fen plant community composition. We identify three indicators of a regime shift occurring in our experimental poor fen system under climate change: nonlinear decline of Sphagnum at temperatures 8 °C above ambient conditions, concomitant increases in Carex spp. at temperatures 4 °C above ambient conditions suggesting a weakening of Sphagnum feedbacks on peat accumulation, and increased variance of the plant community composition and pore water pH through time. A temperature increase of +4 °C appeared to be a threshold for increased vascular plant abundance; however the magnitude of change was species dependent. Elevated temperature combined with elevated CO₂ had a synergistic effect on large graminoid species abundance, with a 15 times increase as compared to control conditions. Community analyses suggested that the balance between dominant plant species was tipped from Sphagnum to a graminoid‐dominated system by the combination of climate change factors. Our findings indicate that changes in peatland plant community composition are likely under future climate change conditions, with a demonstrated shift toward a dominance of graminoid species in poor fens.     

Interactive effects of fertilization and disturbance on community structure and resource availability in an old-field plant community

Summary The interactive effects of fertilization and disturbance on plant community structure and resource availability were studied by supplying four levels of nitrogen and applying four intensities of tilling to a 30 year old field in a factorial design for 2 year. Live above-ground biomass, root biomass, and litter generally increased with nitrogen supply and decreased with disturbance. Species composition varied significantly, with annuals increasing with both nitrogen and disturbance, but with perennials unaffected by nitrogen and decreased by disturbance. Species diversity decreased with disturbance, but decreased with nitrogen only in undisturbed vegetation. Root: shoot ratios decreased with added nitrogen, leaf allocation decreased with disturbance, and flowering allocation increased. Surprisingly, stem allocation was unaffected by disturbance. This result reflected a shift from vertical stems to horizontal stems as disturbance increased. Resource measurements suggested that the vegetation responded to interactions between the treatments as well as to direct treatment effects. Variation in light penetration was reduced by fertilization in undisturbed vegetation but not in tilled plots; variability was not directly affected by disturbance. The availability of nitrogen, the limiting soil nutrient, increased with fertilization but was not significantly affected by disturbance. In contrast, the ratio of ammonium to nitrate was significantly reduced by disturbance but unaffected by supply rates, suggesting that nitrogen may have had different effects under different disturbance regimes, even though its total availability was constant. While many community responses to fertilization and disturbance conformed to those reported earlier, resource and allocation measurements indicated that their interactions are not always predictable from their separate effects.     

The assembly and importance of a novel ecosystem: The ant community of coffee farms in Puerto Rico

Agricultural ecosystems are by their very nature novel and by definition the more general biodiversity associated with them must likewise constitute a novel community. Here, we examine the community of arboreally foraging ants in the coffee agroecosystem of Puerto Rico. We surveyed 20 coffee plants in 25 farms three times in a period of one year. We also conducted a more spatially explicit sampling in two of the farms and conducted a species interaction study between the two most abundant species, Wasmannia auropunctata and Solenopsis invicta, in the laboratory. We find that the majority of the most common species are well‐known invasive ants and that there is a highly variable pattern of dominance that varies considerably over the main coffee producing region of Puerto Rico, suggesting an unusual modality of community structure. The distribution pattern of the two most common species, W. auropunctata and S. invicta, suggests strong competitive exclusion. However, they also have opposite relationships with the percent of shade cover, with W. auropunctata showing a positive relationship with shade, while S. invicta has a negative relationship. The spatial distribution of these two dominant species in the two more intensively studied farms suggests that young colonies of S. invicta can displace W. auropunctata. Laboratory experiments confirm this. In addition to the elaboration of the nature and extent of this novel ant community, we speculate on the possibilities of its active inclusion as part of a biological control system dealing with several coffee pests, including one of the ants itself, W. auropunctata.     

Resource availability and ecosystem size predict food-chain length in pond ecosystems

Food-chain length (FCL) in ecosystems has been studied extensively, and numerous hypotheses to predict FCL, productivity, ecosystem size, and productive space have been proposed. For example, the productivity hypothesis suggests that resource availability limits FCL, whereas the productive-space hypothesis predicts that per-unit-size resource availability and ecosystem size equally limit FCL. However, previous studies have only measured total productivity to test FCL and have never tested the impact of resource availability within a system on FCL. Therefore, we estimated FCL in 15 ponds using stable isotope techniques to test the most common hypotheses for predicting FCL. We also measured total productivity, pond volume (ecosystem size), and edible microalgal carbon (resource availability) in each pond. We found that productive-space (edible carbon+pond volume) was the best model to predict FCL, and FCL was significantly correlated with edible carbon and pond volume. However, factors such as total productivity did not directly predict FCL of the ponds. Therefore, our results suggest that both resource availability and ecosystem size predict FCL in pond ecosystems and play significant roles in maintaining longer FCLs. Furthermore, the productive-space hypothesis appears to be particularly important for determining the FCL of ponds.     

Patterns in the assembly of an island plant community

Aim  To test for patterns in the assembly of an island plant community.
Location  Islands off the west coast of Vancouver Island, British Columbia, Canada.
Methods  Twenty-seven islands were visited by boat, and the abundance of six woody angiosperm species was quantified. Null models were then used to test whether: (1) some species co-occur less than expected by chance (i.e. co-occurrence assembly rule), (2) the incidence and abundance of some species are inversely related to the abundance of other species (i.e. incidence assembly rule), and (3) support for assembly rules precludes evidence for nestedness, which refers to a pattern in species composition in which the species present on depauperate islands form regular subsets of those occurring on progressively more diverse islands.
Results  Most species co-occurred with other species at frequencies expected by chance. However, one species ( Sambucus racemosa ) co-occurred with other species less frequently than randomized expectations. The observed incidence and abundance patterns of most species were also consistent with randomized patterns. However, the incidence and abundance of S. racemosa declined with the abundance of other plant species. Weak, variable support was found for nestedness of the total plant community. However, stronger, consistent support was found after removing S. racemosa from the matrix prior to analyses.
Main conclusions  Most species were assembled on islands in a manner consistent with randomized expectations. However, non-random distributional patterns were observed in one species whose distribution was consistent with the hypothesis that competition limits the assembly of island communities.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.     

植物群落构建机制研究进展

群落构建研究对于解释物种共存和物种多样性的维持是至关重要的,因此一直是生态学研究的中心论题。尽管近年来关于生态位和中性理论的验证研究已经取得了显著的成果,但对于局域群落构建机制的认识仍存在很大争议。随着统计和理论上的进步使得用功能性状和群落谱系结构解释群落构建机制变为可能,主要是通过验证共存物种的性状和谱系距离分布模式来实现。然而,谱系和功能性状不能相互替代,多种生物和非生物因子同时控制着群落构建,基于中性理论的扩散限制、基于生态位的环境过滤和竞争排斥等多个过程可能同时影响着群落的构建。所以,综合考虑多种方法和影响因素探讨植物群落的构建机制,对于预测和解释植被对干扰的响应,理解生物多样性维持机制有重要意义。试图在简要回顾群落构建理论及研究方法发展的基础上,梳理其最新研究进展,并探讨整合功能性状及群落谱系结构的研究方法,解释群落构建和物种多样性维持机制的可能途径。在结合功能性状和谱系结构研究群落构建时,除了考虑空间尺度、环境因子、植被类型外,还应该关注时间尺度、选择性状的种类和数量、性状的种内变异、以及人为干扰等因素对群落构建的影响。     

Resource availability affects the structure of a natural bacteria-bacteriophage community

Antagonistic networks are known to be structured in the wild, but knowledge on how this structure may change as a response to environmental perturbations is scarce. We describe a natural bipartite network between bacteria and lytic bacteriophages, and investigate how it is affected by environmental productivity in the form of different resource levels for the bacteria. We report that low amounts of resource decrease phage generality and lead to less robust and less stable communities. We discuss how resource levels in nature may alter the structure of complex communities.     

Resource availability, species composition and sown density effects on productivity of experimental plant communities

Productivity of artificial grassland communities was investigated in a 6-year field experiment on the Qinghai-Tibetan Plateau. In the experiment, assemblages varying in seven species compositions and four density gradients were grown in fertilized and non-fertilized subplots. We measured biomass of sown species as an indicator of community productivity. In general, 6-years of experiments indicated that: (i) species composition had a significant influence on community productivity. During the initial phase of the experiment, sown density significantly affected community productivity, but the effects disappear with the increase of grown years. This productivity increased with biodiversity increase and fertilization, while the biodiversity effects disappeared when the influence of composition was removed. (ii) The increase of community productivity with biodiversity was resulted from joint effects of selection and complementarity. (iii) With an increase of growth time, the selection effects become weaker while complementarities become enhanced. Influence of density on both effects was significantly different in early stages, but ultimately this all became insignificant. Fertilization dramatically increased the complementarity effects in all experiment processes, but had different influences on selection effects during different experimental period.