Linking individual response to biotic interactions with community structure: a trait-based framework

1.  Due to species-specificity of the outcomes of biotic interactions, it is difficult to generalize from observed biotic interactions at the individual plant level to the effect of those interactions at the community level. To evaluate the importance of biotic interactions in shaping plant communities, it is necessary to understand how the outcomes of the complex interactions observed at the individual level can influence community structure.
2.  Here, we propose a trait-based framework that identifies and organises mechanisms affecting community structure (here described with relative abundances of plant functional traits – i.e. the distribution of trait values at the community level). We applied our approach to a single leaf trait, specific leaf area (SLA), to link individual responses to plant interactions with community structure (SLA distribution observed at the community level) and to test whether biotic interactions can predict the functional composition of subalpine grasslands. We evaluated the generality of our model through a cross-validation with a set of eight subalpine grasslands independent from the four fields used to build the model.
3.  We found that competition and facilitation were able to explain the functional composition of subalpine grasslands, and the relevant fitness components (survival or growth) explaining this link changed depending on the limiting resources. When soil water availability was limiting, positive plant-plant interactions acting on survival were able to explain community structure. In contrast, when no water limitation was observed competition acting on individual growth was the main driver of community structure.
4.  Our framework enables evaluation of the consequences of biotic interactions observed at individual level on community structure, thereby indicating when and where different types of plant-plant interactions are important.     

Truncation of thermal tolerance niches among Australian plants

Aim

Despite recognition that realized distributions inherently underestimate species' physiological tolerances, we are yet to identify the extent of these differences within diverse taxonomic groups. The degree to which species could tolerate environmental conditions outside their observed distributions may have a significant impact on the perceived extinction risk in ecological models. More information on this potential error is required to improve our confidence in management strategies.

Location

Australia.

Time Period

1983–2012.

Major Taxa Studied

Plants.

Methods

To quantify the scale and spatial patterns of this disparity, we estimated the existing tolerance to thermal extremes of 7,124 Australian plants, more than one‐third of the native continental flora, using data from cultivated records at 128 botanical gardens and nurseries. Hierarchical Bayesian beta regression was used to assess whether factors such as realized niches, traits or phylogeny could predict the incidence or magnitude of niche truncation (underestimation of thermal tolerances), while controlling for sources of collection bias.

Results

Approximately half of the cultivated species analysed could tolerate temperature extremes beyond those experienced in their native range. Niche truncation was predictable from the breadth and extremes of their realized niches and by traits such as plant growth form. Phylogenetic relationships with niche truncation were weak and appeared more suited to predicting thermal tolerances directly.

Main conclusions

This study highlights a widespread disparity between realized and potential thermal limits that may have significant implications for species' capacity to persist in situ with a changing climate. Identifying whether thermal niche truncation is the result of biotic interactions, dispersal constraints or other environmental factors could provide significant insight into community assembly at macroecological scales. Estimating niche truncation may help to explain why certain ecological communities are more resilient to change and may potentially improve the reliability of model projections under climate change.     

Nurse plants vs. nurse objects: effects of woody plants and rocky cavities on the recruitment of the Pilosocereus leucocephalus columnar cactus

BACKGROUND AND AIMS: Most studies on cactus recruitment have focused on the role of woody plants as seedling facilitators. Although the spatial association of cacti with objects had been described, the mechanisms underlying this association remain unknown. The aims of this study were to identify which mechanisms facilitate the establishment of a columnar cactus under the shade and protection of objects and to compare these mechanisms with those involved in plant-plant facilitation. METHODS: Three split-split-plot field experiments were conducted to compare the effects of two microhabitats (inside rocky cavities and beneath plant canopies) on seed removal, germination, seedling survivorship and dry weight. Flat, open spaces were used as the control. For each microhabitat, the effect of seed or seedling protection and substrate limitation were explored; aboveground microclimate and some soil properties were also characterized. KEY RESULTS: The permanence of superficial seeds was greater inside rocky cavities than beneath woody plant canopies or on flat, open areas. Germination was similar in cavities and beneath plant canopies, but significantly higher than on flat, open areas. Seedling survivorship was greater beneath plant canopies than inside cavities or on flat, open spaces. CONCLUSIONS: The mechanisms of plant facilitation are different from those of object facilitation. There are seed-seedling conflicts involved in the recruitment of P. leucocephalus: nurse plants favour mainly seedling survivorship by providing a suitable microenvironment, while nurse objects mainly favour seed permanence, by protecting them from predators.     

Plant invaders and their novel natural enemies: who is naïve?

Introduced exotic species encounter a wide range of non‐coevolved enemies and competitors in their new range. Evolutionary novelty is a key aspect of these interactions, but who benefits from novelty: the exotic species or their new antagonists? Paradoxically, the novelty argument has been used to explain both the release from and the suppression by natural enemies. We argue that this paradox can be solved by considering underlying interaction mechanisms. Using plant defenses as a model, we argue that mismatches between plant and enemy interaction traits can enhance plant invasiveness in the case of toxin‐based defenses, whereas invasiveness is counteracted by mismatches in recognition‐based defenses and selective foraging of generalist herbivores on plants with rare toxins. We propose that a mechanistic understanding of ecological mismatches can help to explain and predict when evolutionary novelty will enhance or suppress exotic plant invasiveness. This knowledge may also enhance our understanding of plant abundance following range expansion, or during species replacements along successional stages.     

Do interactions between plant and soil biota change with elevation? A study on Fagus sylvatica

Theoretical models predict weakening of negative biotic interactions and strengthening of positive interactions with increasing abiotic stress. However, most empirical tests have been restricted to plant-plant interactions. No empirical study has examined theoretical predictions of interactions between plants and below-ground micro-organisms, although soil biota strongly regulates plant community composition and dynamics. We examined variability in soil biota effects on tree regeneration across an abiotic gradient. Our candidate tree species was European beech (Fagus sylvatica L.), whose regeneration is extremely responsive to soil biota activity. In a greenhouse experiment, we measured tree survival in sterilized and non-sterilized soils collected across an elevation gradient in the French Alps. Negative effects of soil biota on tree survival decreased with elevation, similar to shifts observed in plant-plant interactions. Hence, soil biota effects must be included in theoretical models of plant biotic interactions to accurately represent and predict the effects of abiotic gradient on plant communities.     

植物间正相互作用对种群动态和群落结构的影响:基于个体模型的研究进展

植物间的相互作用对种群动态和群落结构有着重要的影响。大量的野外实验已经揭示了正相互作用(互利)在群落中的普遍存在及其重要性。为了弥补野外实验方法的不足, 模型方法被越来越多地应用于正相互作用及其生态学效应的研究中。该文基于个体模型研究, 探讨了植物间正相互作用对种群动态和群落结构的影响。介绍了植物间正相互作用的定义和发生机制、植物间相互作用与环境梯度的关系。正相互作用是指发生在相邻的植物个体之间, 至少对其中一个个体有益的相互作用。植物通过直接(生境改善或资源富集)或间接(协同防御等)作用使局部环境有利于邻体而发生正相互作用。胁迫梯度假说认为互利的强度或重要性随着环境胁迫度的增加而增加, 但是越来越多的经验研究认为胁迫梯度假说需要改进。以网格模型和影响域模型为例, 介绍了基于个体的植物间相互作用模型方法。基于个体模型, 对近年来国内外正相互作用对种群时间动态(如生物量-密度关系)、空间分布格局和群落结构(如群落生物量-物种丰富度关系)影响的研究进行了总结。指出未来的研究应集中在对正相互作用概念和机制的理解, 新的模型, 新的种群、群落, 甚至生态系统问题, 以及在全球变化背景下进行相关的研究。     

Community assembly processes restrict the capacity for genetic adaptation under climate change

Given the magnitude and rate of ongoing climate change, the physiological capacity of species to tolerate extreme conditions will play a key role in influencing outcomes for biodiversity. It is also possible that species will respond to changes in climate by shifting their physiological tolerances, through genetic adaptation. How these processes influence biodiversity outcomes will be crucial in determining the most suitable management responses to retain diversity into the future. Here we assess how accounting for physiological tolerances, genetic adaptation and community assembly processes such as species replacement, influence projected climate change outcomes for the flora of Tasmania (all 2051 plant species). We incorporate these processes into the M‐SET metacommunity model and compare four different assumptions of species niches: realized niches, broader physiological tolerances and low or high capacity for genetic adaptation. Accounting for physiological tolerances rather than realized niches had the largest impact on projected outcomes, with 358 fewer species extinctions in the hottest climate scenario (mean = 30 extinctions). In contrast, adding the capacity for species physiological tolerances to shift through genetic adaptation resulted in little additional benefits for biodiversity outcomes, even under an optimistic level of adaptive capacity. We find that this is due largely to community assembly processes such as species replacement restricting the ability of species to persist and adapt in situ, as has been suggested from theoretical metacommunity models applied in simple artificial settings. Our results highlight the importance of accounting for species physiological tolerances and community‐level processes in biodiversity projections, while the potential role for genetic adaptation may be small, requiring further exploration in alternative contexts.     

Plant allelochemical interference or soil chemical ecology?

While allelopathy has been defined as plant-plant chemical interference, there has been much confusion about what the concept encompasses and how important it is in nature. We distinguish between (1) direct plant-plant interference mediated by allelochemicals, and (2) the effects of secondary compounds released by plants on abiotic and biotic soil processes that affect other plants.It very difficult to demonstrate direct effects of chemicals released by a plant on nearby plants. Although soil ecology-mediated effects of secondary plant compounds do not fit the classical concept of allelopathy, we find support in the literature for the hypothesis that the most important effects of compounds released into the soil environment by plants on other plants occur through such indirect effects. The emphasis on, and skepticism of, direct plant-plant allelopathic interference has led some researchers to demand unreasonably high standards of evidence for establishing even the existence of allelopathic interactions, standards that are not demanded for other plant-plant interactions such as resource competition. While the complete elucidation of the mechanisms by which allelochemicals function in the field is many years away, such elucidation is not necessary to establish the existence of allelopathic interactions.We propose that most of the phenomena broadly referred to as allelopathic interference are better conceptualized and investigated in terms of soil chemical ecology. Even when direct plant-plant allelochemical interference occur, the levels of allelochemicals in the environment and their effects on plants are heavily influenced by abiotic and biotic components of the soil ecosystem. Putting allelopathy in the context of soil ecology can further research and reduce some of the less fruitful controversy surrounding the phenomenon.     

植物间挥发物信号的研究进展

植物VOCs信号是植物间进行信息交流的“语言”, 可由多种生物和非生物因素诱导产生。非寄生植物释放的VOCs信号可影响其所在群落中其它植物的种子萌发与幼苗生长; 而寄主植物释放的VOCs信号却是诱导寄生植物种子萌发和幼苗生长的信号物质。VOCs作为植物间的伤害信息可以诱导临近的同种或异种植物做好防御准备, 从而通过主动或间接防御以减少外界的伤害。植物间通过VOCs信号进行信息交流, 从而实现其繁衍与防御。该文通过对VOCs信号的种类、诱导产生因素、传递及作用进行综述, 以期对VOCs信号的研究有所帮助。     

Phylogenetic plant community structure along elevation is lineage specific

The trend of closely related taxa to retain similar environmental preferences mediated by inherited traits suggests that several patterns observed at the community scale originate from longer evolutionary processes. While the effects of phylogenetic relatedness have been previously studied within a single genus or family, lineage‐specific effects on the ecological processes governing community assembly have rarely been studied for entire communities or flora. Here, we measured how community phylogenetic structure varies across a wide elevation gradient for plant lineages represented by 35 families, using a co‐occurrence index and net relatedness index (NRI). We propose a framework that analyses each lineage separately and reveals the trend of ecological assembly at tree nodes. We found prevailing phylogenetic clustering for more ancient nodes and overdispersion in more recent tree nodes. Closely related species may thus rapidly evolve new environmental tolerances to radiate into distinct communities, while older lineages likely retain inherent environmental tolerances to occupy communities in similar environments, either through efficient dispersal mechanisms or the exclusion of older lineages with more divergent environmental tolerances. Our study illustrates the importance of disentangling the patterns of community assembly among lineages to better interpret the ecological role of traits. It also sheds light on studies reporting absence of phylogenetic signal, and opens new perspectives on the analysis of niche and trait conservatism across lineages.     

Multiple effects of shrubs on annual plant communities in arid lands of South Australia

Abstract The presence of shrubs in arid lands creates spatial heterogeneity that affects the distribution and performance of annual plants; several possible mechanisms have been implicated. A preliminary survey in a chenopod shrubland in South Australia showed differences in the distribution of annual plants under canopies of Atriplex vesicaria and Maireana sedifolia (the two dominant shrub species) and open spaces. A series of experiments were conducted to test the potential contribution to these patterns of nutrient enrichment under shrubs, differential seed accumulation, stress reduction by the canopy, competition by shrub roots, and protection against grazing. The germinable soil seed‐bank under A. vesicaria and M. sedifolia was different from that in open spaces, but these differences can only explain a fraction of the differences observed in the growing annual plant community in different microsites. The soil under A. vesicaria had higher total nitrogen content than soil in open spaces, whereas soil under M. sedifolia had lower available phosphorus than open spaces. Although annual plant densities under A. vesicaria were higher than in open spaces, experimental removal of shrubs increased their density, suggesting that shrub canopies inhibit annual plants in this system. Surprisingly, trenching of open areas close to shrubs (severing lateral shrub roots) decreased annual plant density. We suggest that water moves laterally through shrub roots, in a process akin to a hydraulic lift, increasing water availability for the annual plants. Exclusion of vertebrate grazers had a stronger effect on annual plant biomass in open spaces than under M. sedifolia, suggesting that this shrub provides shelter against herbivory. Overall our results show that shrubs can have simultaneously facilitative and inhibitory effects on the annual plant community through different mechanisms, but more importantly that different shrub species have different effects. This is a potential mechanism allowing for species coexistence of annual plants.     

Absence of phylogenetic signal in the niche structure of meadow plant communities

A significant proportion of the global diversity of flowering plants has evolved in recent geological time, probably through adaptive radiation into new niches. However, rapid evolution is at odds with recent research which has suggested that plant ecological traits, including the beta- (or habitat) niche, evolve only slowly. We have quantified traits that determine within-habitat alpha diversity (alpha niches) in two communities in which species segregate on hydrological gradients. Molecular phylogenetic analysis of these data shows practically no evidence of a correlation between the ecological and evolutionary distances separating species, indicating that hydrological alpha niches are evolutionarily labile. We propose that contrasting patterns of evolutionary conservatism for alpha- and beta-niches is a general phenomenon necessitated by the hierarchical filtering of species during community assembly. This determines that species must have similar beta niches in order to occupy the same habitat, but different alpha niches in order to coexist.     

Details of local dispersion improve the fit of neighborhood competition models

I performed a series of greenhouse experiments to explore how patterns in the dispersion of local competitors affect the reproductive performance of Capsella bursa-pastoris, Poa annua and Senecio vulgaris. I manipulated the density and relative frequency of competitors in each of three concentric rings surrounding a central plant, thereby creating a variety patterns by which local competitors were distributed. Neighborhood competition models were used to predict the seed output of these central plants. For both Senecio vulgaris and Capsella bursa-pastoris, I found that models which incorporated the dispersion of competitors, as well as the relative emergence date of plants, performed substantially better than those that considered only the distance between the central plant and each of its competitors. I was unable to measure the seed output of Poa annua but neither emergence data nor the dispersion of competitors were important in determining its final dry-weight. Present address: Department of Biology, Box 1137, Washington University, 1 Brookings Drive, St. Louis, MO 63130, USA     

丛枝菌根菌丝网络在植物互作中的作用机制研究进展

植物间的相互作用是生态学领域关注的重要方向之一,其对高效利用养分资源、提高生产力以及构建植物群落均具有重要意义。丛枝菌根真菌是重要的植物互惠共生微生物,其菌丝可以将邻近植物的根部连接起来,形成共同的菌丝网络(CMNs),这些网络对转移养分、水分以及调节植物群落具有重要作用。近期的研究表明,CMNs可以充当植物之间传递病害和蚜虫诱导信号的通道,并激活邻近植物(未受感染)的化学防御。本文围绕最新的CMNs在植物相互作用中的研究成果,总结了影响CMNs规模及其功能活性的主要因素,阐述了CMNs在植物间养分、水分转移及再分配中的作用,并对CMNs在地下化学信号交流、幼苗建植及群落构建的作用机制进行了系统回顾,最后展望了该研究领域中存在的问题,旨在为进一步理解CMNs在植物互作中的生态学功能提供参考。     

Identifying and evaluating causes of alternative community states in wetland plant communities

Ecological communities can vary greatly in species composition. Often this variation is discontinuous, in that abrupt changes in composition occur over small distances in space or short periods of time. A wide range of hypotheses from different subfields of ecology have been proposed to explain these patterns. I suggest a framework to quantitatively evaluate these hypotheses with observational data by characterizing 1) how community composition varies across sites in space, 2) how community composition varies through time, and 3) the possible drivers of this variation. I applied this approach to understand the community composition of producers in temporary and semipermanent wetlands in Michigan, USA. I identified several distinct community states which were variously dominated by particular plant functional groups (submerged, floating or emergent plants) or had no plants throughout a season. Evaluating possible hypotheses to explain this variation, I found that similar communities were not necessarily clustered near each other, suggesting that dispersal was not limited for these plants. Some sites exhibited a great deal of change in plant composition among years, shifting between two community states, but there was relatively little change at sites within a year. Moreover, these shifts did not occur in a particular order to suggest directional change or repeating cycles. Community composition was associated with several environmental variables such as pH, light and depth, and multivariate analyses suggested that species had complex, nonlinear responses to these possible drivers. Alternative stable states and interactions among multiple nonlinear drivers best explained the patterns observed in these wetlands. By formalizing initial data collection in other systems with the framework suggested here, we may gain insight into the causes of alternative community states beyond wetlands and the role of climate change and other anthropogenic forces in precipitating transitions between states.     

Ecological differentiation among key plant mutualists from a cryptic ant guild

As key dispersers of herbaceous seeds, Aphaenogaster ants strongly influence the distribution of woodland plants in eastern North America. Ants within this genus are difficult to distinguish and often are identified by subgroup, but emerging research suggests they occupy species-specific ecological niches. As such, distinct climatic requirements among Aphaenogaster spp. might result in transient plant interactions with climate change. We examine whether there are ecological and distributional differences among Aphaenogaster species that coincide with current taxonomic differentiations. We use occurrence records for six Aphaenogaster spp. that occur in deciduous forests in eastern North America. We associate the geographic patterning of species occurrence with temperature and precipitation data, and we examine whether unique climatic niches characterize each species. We then predict habitat suitability throughout eastern North America using species distribution models. For verification, we test how well the predicted ranges fit observed occurrences using novel data sets for each species. We find that Aphaenogaster species within this cryptic genus demonstrate unique ecological and geographic signatures. Each species within the subgroup generally responds differently to temperature, and somewhat differently to precipitation and seasonal variance, suggesting unique ecological niches for each species. Our results indicate that each ant species may respond uniquely to changes in climate. Such shifts could disrupt current community associations and biotic interactions with ant-dispersed plants.     

Emergent neutrality or hidden niches?

Emergent neutrality is the idea in community ecology that species interactions may drive a system in a direction where some species become so similar that this similarity will be the primary cause for their coexistence instead of niche differentiation. A recent, widely cited model of emergent neutrality is by Scheffer and van Nes, later applied to species abundance distribution patterns by Vergnon et al. We take issue with the ecological interpretation of this model, demonstrating that it in fact presupposes important differences between superficially similar‐looking species. We argue that the temptation to interpret the model as one of emergent neutrality stems from the fact that these differences are unmodeled and therefore hidden, obscuring the underlying coexistence mechanisms. We therefore claim that the model is actually one of hidden niches, and present several alternative ways to make its hidden portions more explicit. These alterations to the model also make its proper interpretation as one of hidden niches more transparent. We also polemize with the claim of Vergnon et al. that multimodality in species abundance distributions is support for their emergent neutrality model: we demonstrate that appropriate stochastic versions of classical resource partitioning or even neutral models can lead to such patterns in a robust way. Observation of these patterns is therefore inconclusive as to the underlying mechanisms that generate them.     

Mechanisms for locating resources in space and time: Impacts on the abundance of insect herbivores

Herbivorous insects have the problem both of locating appropriate host plants and ensuring that the plant‐feeding stages of their life cycles are synchronized with the times when those hosts provide a high‐quality food resource. Because the taxonomic range of potential hosts is generally narrow, and the temporal window when those hosts are suitable is often relatively short, developmental (especially diapause) and dispersal mechanisms may be critical factors in determining whether or not a species population is successful in a particular plant community. The present paper considers the impact of diapause and dispersal mechanisms on the ability of insect herbivores to cope with two attributes of their host plants: (i) the diversity of the plant community within which the hosts are located; and (ii) the seasonal predictability of host suitability. Some common dispersal mechanisms used by insect herbivores are much more appropriate to low‐diversity than to high‐diversity plant communities and, similarly, some diapause cues are appropriate only to highly predictable plant phenology. Both agriculture and silviculture characteristically manipulate both these attributes of plant communities, that is, in order to make the human use of plants more efficient, cultivation strategies normally both reduce plant species diversity (often to a condition approaching monoculture) and increase the predictability of plant developmental patterns. Consequently, major pest species in managed systems may not be those that are most common in natural systems, and may be difficult to predict in advance.     

Processes underlying the effect of mycorrhizal symbiosis on plant-plant interactions

Mycorrhizal symbiosis has important implications for the diversity and productivity of plant communities. However, our mechanistic understanding of its influence on the outcome of plant-plant interactions is still expanding. In this review we propose a framework that might be useful to efficiently approach the effects of mycorrhizal fungi (MF) on plant-plant interactions. We propose several scenarios that can theoretically result in different outcomes of plant-plant interactions based on the combination of two processes: the diversity of resources provided by MF taxa to their host (resource dissimilarity) and contrasting ways of distributing those resources (resource distribution). Then, we illustrate our arguments with different ecological contexts where certain combinations of these two processes are prone to occur. The proposed framework suggests testable hypotheses that can contribute to elucidate relevant processes underlying the effects of mycorrhizal symbiosis on plant-plant interactions.