Post-fire restoration of Mediterranean forests: Testing assembly rules mediated by facilitation

In view of the importance of facilitative interactions between plants, nurse-based planting has been proposed as a restoration technique for Mediterranean vegetation. However, facilitation efficiency is known to depend on the environmental context and the particular pair of interacting species. Understanding these context- and pair-specific dependences is fundamental to understanding Mediterranean vegetation dynamics and to improving the use of nurse-based plantation for restoration. We assessed the effectiveness of nurse-based plantation and the significance for post-fire restoration of some assembly rules mediated by facilitation. In two nearby areas of different burning ages, we compared seedling establishment of 13 tall shrubs and trees planted in open ground and under nurses. Nurses and planted seedlings were selected from different life-forms. Tests of the assembly rules were provided by the partitioning of the statistical interaction effect between nurse and planted seedling life-forms in a two-factor design. Nurse-based plantation increased seedling survival 2–9 times compared to plantation in open ground, depending on the year. The higher efficiency of nurse-based plantation was consistent for the two burned areas and occurred in many species even in years with contrasting rainfall. We detected pair-specific differences in the efficiency of facilitation. This pair-specificity was partly explained by the dependence between life-forms of nurse and guest species, suggesting the existence of assembly rules. Our results confirm that nurse-based plantation would be an appropriate post-fire restoration technique in Mediterranean mountains under dry-subhumid climate, but suggest that attention to the life-form assemblage rules is needed for an efficient implementation of such technique.     

Functional dissimilarity,not phylogenetic relatedness,determines interspecific interactions among plants in the Tibetan alpine meadows

The hypotheses suggesting that the nature and strength of species interactions should be determined by phylogenetic relatedness have important implications for the understanding of community structure. However, to date, there is limited empirical evidence to support them. At least two basic conditions need to be met in order to expect species interactions to be determined by evolutionary relatedness: a phylogenetic signal in the traits involved in the interactions and changes in the interactions as species are more ecologically similar. Here, we report results of a removal experiment in the Chinese Tibetan plateau in which we directly assessed if the nature and/or strength of interactions among twelve alpine meadow plant species were influenced by their phylogenetic relatedness and/or their functional dissimilarity. For each plant species, we compared its biomass production when grown alone to its biomass in presence of another species and used it as a measure of species interactions. Competition between pairs of species was more frequent than facilitation, with 60% of interactions resulting in plants producing less biomass when a second species was present. We found no effect of phylogenetic relatedness on the prevalence or intensity of competition or facilitation, presumably as none of the studied traits showed phylogenetic signal. Functional dissimilarity based on maximum plant height alone was the best predictor of both the prevalence and strength of competition and facilitation, followed by functional dissimilarity using all five functional traits. Our results pinpoint the limited capacity of phylogenetic relatedness as predictor of species interactions; underlining the limitations of using phylogenetic dispersion patterns to infer mechanisms of community assembly. On the contrary, when the right functional traits are used, functional dissimilarity among species can predict both the nature and strength of their interactions; accentuating the relevance of trait‐based approaches in community ecology research.     

Release from competition and protection determine the outcome of plant interactions along a grazing gradient

Plant interactions are suggested to shift from competition to facilitation and collapse with increasing grazing pressure. The existence of this full range of plant interactions and the role of underlying mechanisms (i.e. release from competition and protecting effect) in response to herbivory remains poorly documented and mainly described in terrestrial systems. We use a large grazing disturbance gradient (five levels of grazing) to test its effect on the outcome of plant interactions and underlying mechanisms in freshwater ecosystems. In a mesocosm experiment, we manipulated the presence of neighbouring plants to test their negative (competition) or protective (facilitation) effects on target plants along the grazing pressure gradient. We predicted that plant interactions 1) shift from competition to indirect facilitation with increased grazing pressure, 2) indirect facilitation collapses at high levels of grazing, 3) release from competition mainly drives the outcome in lowly grazed conditions and, 4) decreased protection occurs in highly grazed conditions responsible for the collapse of facilitation. This study shows the occurrence of the full range of outcomes in plant interactions under a wide spectrum of grazing pressure and indicates how the complex combination of underlying mechanisms shapes variations in plant interactions. We show that both, the release from competition and the increased protection by neighbouring plants drove the shift from competition to indirect facilitation. Declined protection by neighbouring plants resulted in a collapse of indirect facilitation for survival under intense herbivory. Our study provides the first experimental evidence of indirect facilitation structuring freshwater ecosystems thereby validating important ecological concepts mainly developed for terrestrial ecosystems.     

Frequency and intensity of facilitation reveal opposing patterns along a stress gradient

Disentangling the different processes structuring ecological communities is a long‐standing challenge. In species‐rich ecosystems, most emphasis has so far been given to environmental filtering and competition processes, while facilitative interactions between species remain insufficiently studied. Here, we propose an analysis framework that not only allows for identifying pairs of facilitating and facilitated species, but also estimates the strength of facilitation and its variation along environmental gradients. Our framework combines the analysis of both co‐occurrence and co‐abundance patterns using a moving window approach along environmental gradients to control for potentially confounding effects of environmental filtering in the co‐abundance analysis. We first validate our new approach against community assembly simulations, and exemplify its potential on a large 1,134 plant community plots dataset. Our results generally show that facilitation intensity was strongest under cold stress, whereas the proportion of facilitating and facilitated species was higher under drought stress. Moreover, the functional distance between individual facilitated species and their facilitating species significantly changed along the temperature–moisture gradient, and seemed to influence facilitation intensity, although no general positive or general negative trend was discernible among species. The main advantages of our robust framework are as follows: It enables detecting facilitating and facilitated species in species‐rich systems, and it allows identifying the directionality and intensity of facilitation in species pairs as well as its variation across long environmental gradients. It thus opens numerous opportunities for incorporating functional (and phylogenetic) information in the analysis of facilitation patterns. Our case study indicated high complexity in facilitative interactions across the stress gradient and revealed new evidence that facilitation, similarly to competition, can operate between functionally similar and dissimilar species. Extending the analyses to other taxa and ecosystems will foster our understanding how complex interspecific interactions promote biodiversity.     

Plant–plant facilitation increases with reduced phylogenetic relatedness along an elevation gradient

Environmental conditions can modify the intensity and sign of ecological interactions. The stress gradient hypothesis (SGH) predicts that facilitation becomes more important than competition under stressful conditions. To properly test this hypothesis, it is necessary to account for all (not a subset of) interactions occurring in the communities and consider that species do not interact at random but following a specific pattern. We aim to assess elevational changes in facilitation, in terms of species richness, frequency and intensity of the interaction as a function of the evolutionary relatedness between nurses and their associated species. We sampled nurse and their facilitated plant species in two 1000–2000 m. elevation gradients in Mediterranean Chile where low temperature imposes a mortality filter on seedlings. We first estimated the relative importance of facilitation as a mechanism adding new species to communities distributed along these gradients. We then tested whether the frequency and intensity of facilitation increases with elevation, taking into account the evolutionary relatedness of the nurse species and the facilitated species. We found that nurses increase the species richness of the community by up to 35%. Facilitative interactions are more frequent than competitive interactions (56% versus 44%) and facilitation intensity increased with elevation for interactions involving distantly related lineages. Our results highlight the importance of including an evolutionary dimension in the study of facilitation to have a clearer picture of the mechanisms enabling species to coexist and survive under stressful conditions. This knowledge is especially relevant to conserve vulnerable and threatened communities facing new climate scenarios, such as those located in Mediterranean-type ecosystems.     

Asymmetric facilitation can reduce size inequality in plant populations resulting in delayed density‐dependent mortality

Size inequality in plant populations is a ubiquitous feature that has received much attention due to ecological and evolutionary implications. The mechanisms driving size inequality were mainly attributed to different modes of competition (symmetric versus asymmetric), while the potential effects of different modes of facilitation (symmetric versus asymmetric) to this pattern have not yet been fully explored. We employed an individual‐based model to explore the relative roles of both competition and facilitation simultaneously along an environmental stress gradient. Special emphasis was given to the assessment of symmetric facilitation (plants receive benefit from each other equally or proportionally to benefactors’ sizes) and asymmetric facilitation (beneficiary plants receive benefits from benefactor plants that are higher than proportional to the benefactors’ size) in altering plant size inequality. We found that independent of the particular mode of competition, symmetric facilitation generally increased size inequality, whereas asymmetric facilitation decreased it. This pattern was consistent along the stress gradient. Because of their different effects on size inequality, symmetric facilitation accelerated self‐thinning, whereas asymmetric facilitation delayed the onset of density‐dependent mortality, promoting survival under intermediate stress conditions. We compared our model predictions with both 1) a previous modelling study focusing on the effect of (symmetric) facilitation on the size inequality, and 2) re‐analysed data from a published experiment generating asymmetric facilitation of plants against enhanced ultraviolet‐B (UV‐B). Whereas our model predictions and the results of the empirical experiment were consistent, we found that previous theoretical results that solely relied on symmetric facilitation need to be re‐adjusted. Our study showed that combinations of different modes of competition and facilitation can alter size inequality in different ways and with important consequences for the onset of density‐dependent mortality during population development. Explicitly considering different modes and mechanisms of interactions (both facilitation and competition) will improve mechanistic understanding in plant ecology.     

The relative contribution of abundance and phylogeny to the structure of plant facilitation networks

The structure of the real ecological networks is determined by multiple factors including neutral processes, the relative abundances of species, and the phylogenetic relationships of the interacting species. Previous efforts directed to analyze the relative contribution of these factors to network structure have not been able to fully incorporate the phylogenetic relationships between the interacting species. This limitation stems from the difficulty of predicting interaction probabilities based on the independent phylogenies of interacting species (e.g. plants and animals). This is not the case for plant facilitation networks, where nurse and facilitated species evolve in a common phylogeny (e.g. spermatophyte phylogeny). Facilitation networks are characterized by both high nestedness and interactions tending to occur between distantly related nurse and facilitated species. We evaluate the relative contribution of phylogeny and species abundance to explain both the frequency of observed interactions as well as the network structure in a real plant facilitation network at Tehuacán Valley (central Mexico). Our results show that the combined effects of phylogeny and species abundance were, by far, the best predictors of both the frequency of the interactions observed in this community and the parameters (nestedness and connectance) defining the network structure. This finding indicates that species interact proportionally to both their phylogenetic distances and abundances simultaneously. In short, the phylogenetic history of species, acting together with other ecological factors, has a pervasive influence in the structure of ecological networks.     

Burial disturbance leads to facilitation among coastal dune plants

There is growing evidence that interactions among plants can be facilitative as well as competitive, but knowledge of how disturbances influence these interactions and how they vary with species diversity is lacking. We manipulated plant density, species diversity (richness), and a burial disturbance in a controlled, complete factorial experiment to test theories about the relationships among species interactions, disturbance, and richness. The hypotheses tested were 1) burial disturbance reduces plant performance at all levels of density and richness, 2) burial disturbance can cause net plant interactions to become more facilitative, and 3) facilitation increases with species richness. Burial decreased plant survival by 60% and biomass by 50%, supporting the hypothesis that burial reduces plant performance. In the control (unburied) treatment, there was no difference in proportion survival or per plant biomass between low and high density plots, meaning that neither competition nor facilitation was detected. In the buried treatment, however, high density plots had significantly greater survival and greater per plant biomass than the low density plots, indicating net facilitative interactions. Thus facilitation occurred in the buried treatment and not in the unburied control plots, supporting the hypothesis that facilitation increases with increasing disturbance severity. The hypothesis that facilitation increases with increasing species richness was not supported. Richness did not affect survival or biomass, and there was no richness by burial treatment interaction, indicating that richness did not influence the response of the community to burial. The influence of the disturbance on plant interactions was thus consistent across levels of richness, increasing the generality of the relationship between disturbance and facilitation.     

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species‐specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species‐specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.     

Increased competition does not lead to increased phylogenetic overdispersion in a native grassland

That competition is stronger among closely related species and leads to phylogenetic overdispersion is a common assumption in community ecology. However, tests of this assumption are rare and field‐based experiments lacking. We tested the relationship between competition, the degree of relatedness, and overdispersion among plants experimentally and using a field survey in a native grassland. Relatedness did not affect competition, nor was competition associated with phylogenetic overdispersion. Further, there was only weak evidence for increased overdispersion at spatial scales where plants are likely to compete. These results challenge traditional theory, but are consistent with recent theories regarding the mechanisms of plant competition and its potential effect on phylogenetic structure. We suggest that specific conditions related to the form of competition and trait conservatism must be met for competition to cause phylogenetic overdispersion. Consequently, overdispersion as a result of competition is likely to be rare in natural communities.     

Plant Facilitation on a Mine Tailings Dump

Facilitation is a potentially useful tool in restoration efforts. We investigated the causes of facilitation in planted Picea mariana seedlings in an unvegetated mine tailings dump. Clusters of plants doubled the survival rate in the first growing season compared to single plants. In the first year mycorrhizal inoculation had no effect on survival, but by the second growing season only mycorrhizal inoculated plants survived, most of these being in plant clusters. This suggests that facilitation in this environment is partly the result of interactions with mycorrhizae.     

How Do Alien Plants Fit in the Space-Phylogeny Matrix?

Recent advances in the field of plant community phylogenetics and invasion phylogenetics are mostly based on plot-level data, which do not take into consideration the spatial arrangement of individual plants within the plot. Here we use within-plot plant coordinates to investigate the link between the physical distance separating plants, and their phylogenetic relatedness. We look at two vegetation types (forest and grassland, similar in species richness and in the proportion of alien invasive plants) in subtropical coastal KwaZulu-Natal, South Africa. The relationship between phylogenetic distance and physical distance is weak in grassland (characterised by higher plant densities and low phylogenetic diversity), and varies substantially in forest vegetation (variable plant density, higher phylogenetic diversity). There is no significant relationship between the proportion of alien plants in the plots and the strength of the physical-phylogenetic distance relationship, suggesting that alien plants are well integrated in the local spatial-phylogenetic landscape.     

植物邻体间的正相互作用

植物间的正负相互作用是构建植被群落的重要因素,也是群落生态学研究的中心内容之一。近20a来,植物间正相互作用的研究得到快速发展。综述了正相互作用的定义,不同植物群落中的直接、间接正相互作用及其发生机制,正相互作用研究的实验和模型方法,正负相互作用随胁迫梯度的变化及正相互作用对群落构建的影响。探讨了正相互作用研究前景:(1)进一步理解正负相互作用的平衡及其对群落构建的影响;(2)加深对全球变暖背景下的正相互作用的认识;(3)需把正相互作用研究同进化联系起来;(4)充分发挥正相互作用在生态系统中的推动力作用,把正相互作用应用到生态恢复中,为恢复退化生态系统服务。     

The importance of conspecific facilitation during recruitment and regeneration: A case study in degraded mangroves

Facilitation can affect positively the survival, growth and fitness of neighboring plants and is able to promote the establishment of plant species under harsh environmental conditions. This study is one of the few to focus on conspecific nurse plants enabling the establishment of tree followers and the generally unknown interplay between facilitation and competition during recruitment and establishment.Field studies were conducted in an hypersaline area in north Brazil where the hydrological regime was disturbed by road construction in 1974. Avicennia germinans shrubs were able to recolonize subareas.Three different stages of re-colonization were identified by satellite imagery based on the vegetation coverage and were defined as the early, middle, and late stages. Different plant parameters, including height and crown radius of individual trees, were measured, and all plants separating mangrove seedlings and larger plants were stem-mapped.The Hegyi index was adapted to measure local interaction intensity, considering both positive (facilitation) and negative (competition) interactions among neighboring plants. Spatial point pattern analyses were combined with the interaction index to obtain a better overview of the strength and importance of the plant interactions within the different recolonizing stages.The spatial patterns displayed aggregation for all plants in all developmental stages. This is supported by the interaction index calculated for seedlings. The index, however, provides an early signal for the switching of plant interactions from facilitation to competition as the developmental stage increased. This feature remains undetected in the spatial patterns because self-thinning processes are linked to individual growth processes and require more time to take place.This study broadens the ecological concept of nurse plants from hetero- to conspecific, including the importance and strength of plant interactions. The studied degraded areas are recolonizing naturally, and conclusions can be drawn for restoration projects, which usually do not take facilitation effects into account.     

Limiting similarity and Darwin’s naturalization hypothesis: understanding the drivers of biotic resistance against invasive plant species

Several hypotheses have been proposed to explain biotic resistance of a recipient plant community based on reduced niche opportunities for invasive alien plant species. The limiting similarity hypothesis predicts that invasive species are less likely to establish in communities of species holding similar functional traits. Likewise, Darwin’s naturalization hypothesis states that invasive species closely related to the native community would be less successful. We tested both using the invasive alien Ambrosia artemisiifolia L. and Solidago gigantea Aiton, and grassland species used for ecological restoration in central Europe. We classified all plant species into groups based on functional traits obtained from trait databases and calculated the phylogenetic distance among them. In a greenhouse experiment, we submitted the two invasive species at two propagule pressures to competition with communities of ten native species from the same functional group. In another experiment, they were submitted to pairwise competition with native species selected from each functional group. At the community level, highest suppression for both invasive species was observed at low propagule pressure and not explained by similarity in functional traits. Moreover, suppression decreased asymptotically with increasing phylogenetic distance to species of the native community. When submitted to pairwise competition, suppression for both invasive species was also better explained by phylogenetic distance. Overall, our results support Darwin’s naturalization hypothesis but not the limiting similarity hypothesis based on the selected traits. Biotic resistance of native communities against invasive species at an early stage of establishment is enhanced by competitive traits and phylogenetic relatedness.     

Phylogenetic distance among beneficiary species in a cushion plant species explains interaction outcome

Determining which drivers lead to a specific species assemblage is a central issue in community ecology. Although many processes are involved, plant–plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both communities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context‐dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly‐related species showed an opposite pattern with environmental severity.     

Does phylogenetic relatedness influence the strength of competition among vascular plants?

A widely assumed but largely untested hypothesis central to ecology and evolutionary biology has been Charles Darwin's suggestion that closely related species will be more ecologically similar, and thus will compete more strongly with each other than they will with more distantly related species. We provide one of the first direct tests of the “competition-relatedness hypothesis” by combining two data sets: the relative competitive ability of 50 vascular plant species competing against 92 competitor species measured in five multi-species experiments, and measures of the phylogenetic relatedness of these species. In contrast to Darwin's assertion, there were weak relationships between the strength of competition and phylogenetic relatedness. Across all species studied, the competition-relatedness relationship was weak and not significant. This overall lack of pattern masked different responses of monocot and eudicot focal (phytometer) species. When monocots served as the focal (phytometer) species, the intensity of competition increased with the phylogenetic distance separating species, while competition decreased with phylogenetic distance for eudicot phytometers. These results were driven by the monocot-eudicot evolutionary split, such that monocots were poor competitors against eudicots, while eudicots are most strongly suppressed by other eudicots. There was no relationship between relatedness and competition for eudicots competing with other eudicots, while monocots did compete more intensely with closely related monocots than with distantly related monocots. Overall, the relationships between competition intensity and relatedness were weak compared to the strong and consistent relationships between competitive ability and functional traits such as plant size that have been reported by other studies. We suggest that Darwin's assertion that competition will be strongest among closely related species is not supported by empirical data, at least for the 142 vascular plant species in this study.     

The influence of warming on the biogeographic and phylogenetic dependence of herbivore–plant interactions

Evolutionary experience and the phylogenetic relationships of plants have both been proposed to influence herbivore–plant interactions and plant invasion success. However, the direction and magnitude of these effects, and how such patterns are altered with increasing temperature, are rarely studied. Through laboratory functional response experiments, we tested whether the per capita feeding efficiency of an invasive generalist herbivore, the golden apple snail, Pomacea canaliculata, is dependent on the biogeographic origin and phylogenetic relatedness of host plants, and how increasing temperature alters these dependencies. The feeding efficiency of the herbivore was highest on plant species with which it had no shared evolutionary history, that is, novel plants. Further, among evolutionarily familiar plants, snail feeding efficiency was higher on those species more closely related to the novel plants. However, these biogeographic dependencies became less pronounced with increasing temperature, whereas the phylogenetic dependence was unaffected. Collectively, our findings indicate that the susceptibility of plants to this invasive herbivore is mediated by both biogeographic origin and phylogenetic relatedness. We hypothesize that warming erodes the influence of evolutionary exposure, thereby altering herbivore–plant interactions and perhaps the invasion success of plants.     

Manifold influences of phylogenetic structure on a plant–herbivore network

Ecologists are increasingly aware of the interplay between evolutionary history and ecological processes in shaping current species interaction patterns. The inclusion of phylogenetic relationships in studies of species interaction networks has shown that closely related species commonly interact with sets of similar species. Notably, the degree of phylogenetic conservatism in antagonistic ecological interactions is frequently stronger among species at lower trophic levels than among those at higher trophic levels. One hypothesis that accounts for this asymmetry is that competition among consumer species promotes resource partitioning and offsets the maintenance of dietary similarity by phylogenetic inertia. Here, we used a regional plant–herbivore network comprised of Asteraceae species and flower‐head endophagous insects to evaluate how the strength of phylogenetic conservatism in species interactions differs between the two trophic levels. We also addressed whether the asymmetry in the strength of the phylogenetic signal between plants and animals depends on the overall degree of relatedness among the herbivores. We show that, beyond the previously reported compositional similarity, closely related species also share a greater proportion of counterpart phylogenetic history, both for resource and consumer species. Comparison of the patterns found in the entire network with those found in subnetworks composed of more phylogenetically restricted groups of herbivores provides evidence that resource partitioning occurs mostly at deeper phylogenetic levels, so that a positive phylogenetic signal in antagonist similarity is detectable even between closely related consumers in monophyletic subnetworks. The asymmetry in signal strength between trophic levels is most apparent in the way network modules reflect resource phylogeny, both for the entire network and for subnetworks. Taken together, these results suggest that evolutionary processes, such as phylogenetic conservatism and independent colonization history of the insect groups may be the main forces generating the phylogenetic structure observed in this particular plant–herbivore network system.     

Positive Interactions： Crucial Organizers in a Plant Community

For more than a century, ecologists have concentrated on competition as a crucial process for community organization. However, more recent experimental investigations have uncovered the striking Influence of positive Interactions on the organization of plant communities. Complex combinations of competition and positive interactions operating simultaneously among plant species seem to be widespread In nature. In the present paper, we reviewed the mechanism and ecological importance of positive Interactions In plant communities, emphasizing the certainties and uncertainties that have made It an attractive area of research. Positive Interactions, or facilitation, occur when one species enhances the survival, growth, or richness of another. The Importance of facilitation in plant organization increases with ablotlc stress and the relative Importance of competition decreases. Only by combining plant interactions and the many fields of biology can we fully understand how and when the positive Interactions occur.