Facilitation and plant phenotypic evolution.

While antagonistic interactions between plants have been a major topic of eco-evolutionary research, little evidence exists on the evolution of positive plant interactions (i.e., plant facilitation). Here, we first summarize the existing empirical evidence on the role of facilitation as a selection pressure on plants. Then, we develop a theoretical eco-evolutionary framework based on fitness-trait functions and interaction effectiveness that provides predictions for how facilitation-related traits may evolve. As evolution may act at levels beyond the individual (such as groups or species), we discuss the subject of the units of evolutionary selection through facilitation. Finally, we use the proposed formal evolutionary framework for facilitation to identify areas of future research based on the knowledge gaps detected.

A nurse plant benefits from facilitative interactions through mycorrhizae.

Plant facilitation promotes coexistence by maintaining differences in the regeneration niche because some nurse species recruit under arid conditions, whereas facilitated species recruit under more mesic conditions. In one Mexican community, 95% of species recruit through facilitation; Mimosa luisana being a keystone nurse for many of them. M. luisana individuals manifest greater fitness when growing in association with their facilitated plants than when growing in isolation. This observation suggests that nurses also benefit from their facilitated plants, a benefit thought to be mediated by mycorrhizal fungi. Under field conditions, we experimentally tested whether mycorrhizal fungi mediate the increased fitness that M. luisana experiences when growing in association with its facilitated plants. We applied fungicide to the soil for nurse plants growing alone and growing in association with their facilitated plants in order to reduce the mycorrhizal colonisation of roots. We then assessed the quantity and quality of seed production of M. luisana in four treatments (isolated-control, isolated-fungicide, associated-control and associated-fungicide). Fungicide application reduced the percentage root length colonised by mycorrhizae and reduced fitness of M. luisana when growing in association with their facilitated plants but not when growing in isolation. This reduction was reflected in the total number of seeds, number of seeds per pod, seed mass and seed viability. These results suggest that nurses benefit from the presence of their facilitated plants through links established by mycorrhizae, indicating that both plants and belowground mutualistic communities are all part of one system, coexisting by means of intrinsically linked interactions.

Fungal phylogenetic diversity drives plant facilitation.

Plant-plant facilitation is a crucial ecological process, as many plant species (facilitated) require the presence of an established individual (nurse) to recruit. Some plant facilitative interactions disappear during the ontogenetic development of the facilitated plant but others persist, even when the two plants are adults. We test whether the persistence of plant facilitative interactions is explained by the phylogenetic diversity of mutualistic and non-mutualistic fungi that the nurse and the facilitated species add to the shared rhizosphere. We classify plant facilitative interactions as persistent and non-persistent interactions and quantify the phylogenetic diversity of mutualistic and non-mutualistic fungi added by the plant species to the shared rhizosphere. Our results show that the facilitated species add less phylogenetic diversity of non-mutualistic fungi when plant facilitative interactions persist than when they do not persist. However, persistent and non-persistent facilitative interactions did not differ in the phylogenetic diversity of mutualistic fungi added by the facilitated species to the shared rhizosphere. Finally, the fungal phylogenetic diversity added by the nurse to the shared rhizosphere did not differ between persistent and non-persistent interactions. This study suggests that considering the fungal associates of the plant species involved in facilitative interactions can shed light on the mechanisms of persistence for plant-plant interactions.

Evidence for phylogenetic correlation of plant-AMF assemblages?

BACKGROUND AND AIMS: Specificity in biotic interactions is mediated' by functional traits inducing shifts in the community species composition. Functional traits are often evolutionarily conserved, resulting in closely related species tending to interact with similar species. This tendency may initially shape the phylogenetic composition of coexisting guilds, but other intraguild ecological processes may either blur or promote the mirroring of the phylogenetic compositions between guilds. The roles of intra- and interguild interactions in shaping the phylogenetic community composition are largely unknown, beyond the mere selectivity in the interguild interactions. Plant facilitation is a phylogenetically structured species-specific process involving interactions not only between the same guild of plants, but also between plants and other guilds such as arbuscular mycorrhizal fungi (AMF). In this study it is hypothesized that reciprocal plant-AMF interactions will leave an interdependent phylogenetic signal in the community composition of both plants and AMF. METHODS: A correlation was used to test for a relationship between the phylogenetic composition of plant and AMF assemblages in a patchy xeric shrubland environment shaped by plant facilitation. In addition, a null model was used to test whether this correlation can be solely explained by selectivity in plant-AMF interactions. KEY RESULTS: A significant correlation was observed between the phylogenetic composition of plant and AMF assemblages. Plant phylogenetic composition in a patch was related to the predominance of plant species with high nursery quality that can influence the community assembly. AMF phylogenetic composition was related to the AMF phylogenetic diversity in each patch. CONCLUSIONS: This study shows that shifts in the phylogenetic composition of plants and AMF assemblages do not occur independently. It is suggested that besides selectivity in plant-AMF interactions, inter-related succession dynamics of plants and AMF within patches could be an ecological mechanism driving community assembly. Future lines of research might explore whether interlinked above- and below-ground dynamics could be occurring across multiple guilds simultaneously.

Facilitation allows plant coexistence in Cuban serpentine soils.

Serpentine soils represent stressful habitats where plants have to cope with heavy metals, moisture limitation and low nutrient availability. We propose that facilitation is an important mechanism structuring plant communities under such stressful conditions. Facilitation has been shown to generate the spatial association of species, forming discrete vegetation patches of phylogenetically distant species. We measured these spatial and phylogenetic signatures left by facilitation in a serpentine plant community of central Cuba. Our results show that seedlings preferentially grow under plants of different species, and that adults are significantly aggregated into vegetation patches. In these patches, adults tend to co-occur with distant relatives, ultimately forming phylogenetically diverse neighbourhoods. We discuss possible mechanisms explaining how species adapted to serpentine areas may be acting as nurses, reducing the stressful conditions for the establishment of other species.

Plant facilitation occurs between species differing in their associated arbuscular mycorrhizal fungi.

Complementary beneficial effects of different arbuscular mycorrhizal fungi (AMF) can result in a more efficient exploitation of the soil nutrients available, thus influencing plant communities. Here, we hypothesize that plant-AMF specificity is mediated by phylogenetic constraints defining possible interactions, and that plant-AMF interaction patterns can influence plant-plant facilitation specificity. We reanalyzed previous data describing plant-plant and plant-AMF interaction at the community level to specifically test for a phylogenetic signal on plant and AMF interactions and for a relationship between plant-plant facilitation specificity and plant species differences in their AMF associates. Closely related AMF operational taxonomical units (OTUs) tend to interact with the same plant species, but there is not a significant signal in the interaction through the plant phylogeny. This indicates that the similarity in the AMF associates of two plant species is independent of their phylogenetic relatedness. Interestingly, plant-AMF interactions match plant facilitation specificity, with pairs of plant species recruiting more frequently under each other tending to have different AMF associates. An increment of AMF diversity in the rhizosphere, as a result of plant-AMF and plant-plant selectivity, is suggested as a potential driver of plant-plant facilitation. This study highlights the role of plant-AMF interactions in shaping plant community assemblages.

The network structure of plant-arbuscular mycorrhizal fungi.

Ecological network theory predicts that in mutualistic systems specialists tend to interact with a subset of species with which generalists interact (i.e. nestedness). Approaching plant-arbuscular mycorrhizal fungi (AMF) association using network analyses will allow the generality of this pattern to be expanded to the ubiquitous plant-AMF mutualism. Based on certain plant-AMF specificity recently suggested, networks are expected to be nested as a result of their mutualistic nature, and modular, with certain species interacting more tightly than others. Network analyses were used to test for nestedness and modularity and to compare the different contribution of plant and AMF to the overall nestedness. Plant-AMF networks share general network properties with other mutualisms. Plant species with few AMFs in their roots tend to associate with those AMFs recorded in most plant species. AMFs present in a few plant species occur in plant species sheltering most AMF (i.e. nestedness). This plant-AMF network presents weakly interlinked subsets of species, strongly connected internally (i.e. modularity). Both plants and AMF show a nested structure, although AMFs have lower nestedness than plants. The plant-AMF interaction pattern is interpreted in the context of how plant-AMF associations can be underlying mechanisms shaping plant community assemblages.

Fire and resprouting in Mediterranean ecosystems: insights from an external biogeographical region, the mexical shrubland.

We investigated modes of regeneration of dominant species of the mexical vegetation after fire. The mexical shrubland shows a remarkable structural, morphological, and floristic similarity to Mediterranean-type vegetation and is considered a relict of the Madro-Tertiary Geoflora under a non-Mediterranean climate. This vegetation provides an ideal scenario to test the role of fire in Mediterranean ecosystems because historical fire occurrence is absent and the species assembly is constituted mostly by Madro-Tertiary elements and Neotropical species (some of them, endemic species from Mexico). The existence of congeneric species of the California chaparral allows us to determine the regeneration ability of these communities after fire in relation to resprouting and seeding strategies, which are widespread modes reported in the Mediterranean-type vegetation. By the experimental application of fire in the two biogeographical groups of species, we tested the hypothesis that low resprouting ability of California congeneric species (Madro-Tertiary species) after fire would indicate that fire has played an important selective force in the resprouting habit. A low resprouting ability in the Neotropical group of species would suggest that fire has molded the set of species dominating fire-prone environments.Our results indicated that resprouting is a widespread trait in the mexical species characterized by the presence of lignotubers and burls. Resprouting can be considered an ancient trait, probably linked to losses of aboveground biomass, that became a pre-adaptation in Mediterranean fire-prone communities. The Neotropical group of species showed less ability to regenerate after fire, and small plants were more likely to die after disturbance in this group than in the Madro-Tertiary group. The resprouting feature and the seeder strategy of other species after a fire in the mexical shrubland are similar to Mediterranean-type ecosystems, emphasizing their common origin and the relevance of phylogenetic and biogeographical studies to explain current patterns of vegetation.

Morphological variation and the process of domestication of Stenocereus stellatus (Cactaceae) in Central Mexico.

Morphological variation was analyzed in wild, managed in situ, and cultivated populations of the columnar cactus Stenocereus stellatus in central Mexico. The purpose was to evaluate whether morphological divergence between manipulated and wild populations has resulted from domestication processes. Variation of 23 morphological characters was analyzed among 324 individuals from 19 populations of the Tehuacán Valley and La Mixteca Baja. Multivariate statistical analyses were used to group individuals and populations according to their morphological similarity. Individuals grouped according to the way of management and fruit characteristics were the most relevant for grouping. Within each region, sweet fruits with pulp colors other than red were more frequent in cultivated populations, where fruits were also larger, contained more and bigger seeds, and had thinner peel and fewer spines than fruits from wild individuals. Phenotypes common in managed in situ and cultivated populations generally occur in the wild but in lower frequencies. Artificial selection has thus operated by enhancing and maintaining desirable rare phenotypes in managed in situ and cultivated populations, causing divergent patterns of morphological variation from wild populations. Cultivation has caused the strongest level of divergence, but divergence has also been significant with management of wild populations in situ.

Reproductive biology and the process of domestication of the columnar cactus Stenocereus Stellatus in Central Mexico.

Pollination biology, breeding system, and floral phenology of the columnar cactus Stenocereus stellatus were studied in wild, wild managed in situ and cultivated populations of central Mexico, in order to examine whether these aspects have been modified under domestication and whether they determine reproductive barriers between wild and manipulated individuals. Individuals of both wild and manipulated populations are self-incompatible, indicating that artificial selection has not modified the breeding system. Their pollination biology is also similar. Anthesis is mainly nocturnal, with a peak of nectar production between 0200 and 0400 when the stigma presents maximum turgidity. Nocturnal visitors are the effective pollinators. Nearly 75% of flowers exposed for nocturnal pollination set fruit, while none of the flowers exposed for diurnal pollination produced fruits. The bats Leptonycteris curasoae, L. nivalis, and Choeronycteris mexicana (Glossophaginae) are the most likely pollinators, and their time of foraging is synchronized with the time of nectar production and stigma receptivity in S. stellatus. Bats potentially move pollen over a considerable distance, so there is apparently no spatial isolation to prevent pollen exchange between wild and cultivated populations. Phenological studies showed that there are also no apparent temporal barriers. However, manual cross pollination failed between some domesticated and wild phenotypes, suggesting that gene flow between wild and cultivated populations might be limited by pollen incompatibility.

The chaparral vegetation in Mexico undernonmediterranean climate: the convergence and Madrean-Tethyan hypothesesreconsidered.

A comparative study between an unburned evergreen sclerophyllousvegetation located in south-central Mexico under a wet-summer climate,with mediterranean regions was conducted in order to re-analyzevegetation and plant characters claimed to converge under mediterraneanclimates. The comparison considered floristic composition,plant-community structure, and plant characters as adaptations tomediterranean climates and analyzed them by means of a correspondenceanalysis, considering a tropical spiny shrubland as the external group.We made a species register of the number of species that resproutedafter a fire occurred in 1995 and a distribution map of the evergreensclerophyllous vegetation in Mexico (mexical) under nonmediterraneanclimates.The Tehuacán mexical does not differ from the evergreensclerophyllous areas of Chile, California, Australia, and theMediterranean Basin, according to a correspondence analysis, whichordinated the Tehuacán mexical closer to the mediterranean areasthan to the external group.All the vegetation and floristic characteristics of the mexical, aswell as its distribution along the rain-shadowed mountain parts ofMexico, support its origin in the Madrean-Tethyan hypothesis of Axelrod.Therefore, these results allow to expand the convergence paradigm of thechaparral under an integrative view, in which a general trend to ariditymight explain floristic and adaptive patterns detected in theseenvironments.