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The diversity of functional and life-history traits of organisms depends on adaptation as well as the legacy of shared ancestry. Although the evolution of traits in macro-organisms is well studied, relatively little is known about character evolution in micro-organisms. Here, we surveyed an ancient and ecologically important group of microbial plant symbionts, the arbuscular mycorrhizal (AM) fungi, and tested hypotheses about the evolution of functional and life-history traits. Variation in the extent of root and soil colonization by AM fungi is constrained to a few nodes basal to the most diverse groups within the phylum, with relatively little variation associated with recent divergences. We found no evidence for a trade-off in biomass allocated to root versus soil colonization in three published glasshouse experiments; rather these traits were positively correlated. Partial support was observed for correlated evolution between fungal colonization strategies and functional benefits of the symbiosis to host plants. The evolution of increased soil colonization was positively correlated with total plant biomass and shoot phosphorus content. Although the effect of AM fungi on infection by root pathogens was phylogenetically conserved, there was no evidence for correlated evolution between the extent of AM fungal root colonization and pathogen infection. Variability in colonization strategies evolved early in the diversification of AM fungi, and we propose that these strategies were influenced by functional interactions with host plants, resulting in an evolutionary stasis resembling trait conservatism.

Summary Plant functional traits have revealed trade‐offs related to life‐history adaptations, geographical distributions, and ecosystem processes. Fine roots are essential in plant resource acquisition and play an important role in soil carbon cycling. Nonetheless, root trait variation is still poorly quantified and rarely related to the rest of the plant. We examined chemical and morphological traits of 34 temperate arbuscular mycorrhizal tree species, representing three main angiosperm clades (super‐orders asterid, magnoliid and rosid). We tested to what extent fine root chemical and morphological traits were correlated similarly to the leaf economical spectrum (LES) or were structured by ancestral affiliations among species. Root traits did not display the same trade‐offs as leaves (e.g. specific root length was not correlated with root N, whereas specific leaf area was correlated with leaf N). Moreover, 75% of below‐ground traits were phylogenetically structured according to Pagel's λ and Abouheif's Cmean autocorrelation tests, as opposed to 28% of above‐ground traits. Magnoliids showed thicker, less branched roots than asterids or rosids, but rosid roots exhibited lower N and higher non‐acid‐hydrolysable (e.g. lignin) content than other species. In contrast, leaf traits did not differ significantly among super‐orders. At the whole‐tree level, chemical traits such as nitrogen tissue content and lignin content were correlated between above and below‐ground organs. The distribution of root traits in woody temperate trees was better explained by shared ancestry than by the nutrient content and structural trade‐offs expected by the LES hypothesis. Root chemistry and morphology differed substantially among species belonging to different super‐orders, suggesting deep divergences in resource acquisition strategies among major angiosperm groups. Although we found partial support for the idea of whole‐plant integration based on corresponding nitrogen content across all organs (i.e. a plant economics spectrum), our study stresses phylogenetic affiliation as the primary driver of root trait distributions among angiosperms, a pattern that could be easily overlooked based solely on above‐ground observations. Lay Summary

The relative importance of biotic, abiotic, and stochastic processes in structuring ecological communities continues to be a central focus in community ecology. In order to assess the role of phylogenetic relatedness on the nature of biodiversity we first quantified the degree of phylogenetic niche conservatism of several plant traits linked to plant form and function. Next we quantified the degree of phylogenetic relatedness across two fundamental scaling dimensions: plant size and neighborhood size. The results show that phylogenetic niche conservatism is likely widespread, indicating that closely related species are more functionally similar than distantly related species. Utilizing this information we show that three of five tropical forest dynamics plots (FDPs) exhibit similar scale-dependent patterns of phylogenetic structuring using only a spatial scaling axis. When spatial- and size-scaling axes were analyzed in concert, phylogenetic overdispersion of co-occurring species was most important at small spatial scales and in four of five FDPs for the largest size class. These results suggest that phylogenetic relatedness is increasingly important: (1) at small spatial scales, where phylogenetic overdispersion is more common, and (2) in large size classes, where phylogenetic overdispersion becomes more common throughout ontogeny. Collectively, our results highlight the critical spatial and size scales at which the degree of phylogenetic relatedness between constituent species influences the structuring of tropical forest diversity.

Since root endophytes may ameliorate drought stress, understanding which plants associate with endophytes is important, especially in arid ecosystems. Here, the root endophytes were characterized of 42 plants from an arid region of Argentina. Colonization by arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSEs) was related to plant functional type (PFT), family, and phylogenetic relatedness. Overall, three main findings were observed. Firstly, only moderate levels of endophyte associations were found across all taxa (e.g., most Poaceae were not colonized by endophytes despite numerous accounts of colonization by AMF and DSEs). We determined 69 % of plant taxa associated with some form of root endophyte but levels were lower than other regional studies. Secondly, comparisons by PFT and phylogeny were often qualitatively similar (e.g., succulents and Portulacineae consistently lacked AMF; variation occurred among terrestrial vs. epiphytic bromeliads) and often differed from comparisons based on plant family. Thirdly, comparisons by plant family often failed to account for important variation either within families (e.g., Bromeliaceae and Poaceae) or trait conservatism among related families (i.e., Rosidae consistently lacked DSEs and Portulacineae lacked AMF). This study indicates the value of comparing numerous taxa based on PFTs and phylogenetic similarity. Overall, the results suggest an uncertain benefit of endophytes in extremely arid environments where plant traits like succulence may obviate the need to establish associations.

Functional traits mediate ecological responses of organisms to the environment, determining community structure. Community-weighted trait means (CWM) are often used to characterize communities by combining information on species traits and distribution. Relating CWM variation to environmental gradients allows for evaluating species sorting across the metacommunity, either based on correlation tests or ordinary least squares (OLS) models. Yet, it is not clear if phylogenetic signal in both traits and species distribution affect those analyses. On one hand, phylogenetic signal might indicate niche conservatism along clade evolution, reinforcing the environmental signal in trait assembly patterns. On the other hand, it might introduce phylogenetic autocorrelation to mean trait variation among communities. Under this latter scenario, phylogenetic signal might inflate type I error in analysis relating CWM variation to environmental gradients. We explore multiple ways phylogenetic history may influence analysis relating CWM to environmental gradients. We propose the concept of neutral trait diffusion, which predicts that for a functional trait x, CWM variation among local communities does not deviate from the expectation that x evolved according to a neutral evolutionary process. Based on this framework we introduce a graphical tool called neutral trait diffusion representation (NTDR) that allows for the evaluation of whether it is necessary to carry out phylogenetic correction in the trait prior to analyzing the association between CWM and environmental gradients. We illustrate the NTDR approach using simulated traits, phylogenies and metacommunities. We show that even under moderate phylogenetic signal in both the trait used to define CWM and species distribution across communities, OLS models relating CWM variation to environmental gradients lead to inflated type I error when testing the null hypothesis of no association between CWM and environmental gradient. To overcome this issue, we propose a phylogenetic correction for OLS models and evaluate its statistical performance (type I error and power). Phylogeny-corrected OLS models successfully control for type I error in analysis relating CWM variation to environmental gradients but may show decreased power. Combining the exploratory tool of NTDR and phylogenetic correction in traits, when necessary, guarantees more precise inferences about the environmental forces driving trait-mediated species sorting across metacommunities.

Phylogenetic niche conservatism (PNC) is the tendency of lineages to retain their niche-related traits through speciation events. A recent surge in the availability of well-sampled molecular phylogenies has stimulated phylogenetic approaches to understanding ecological processes at large geographical scales and through macroevolutionary time. We stress that PNC is a pattern, not a process, and is found only in some traits and some lineages. At the simplest level, a pattern of PNC is an inevitable consequence of evolution – descent with modification and divergence of lineages – but several intrinsic causes, including physicochemical, developmental and genetic constraints, can lead directly to a marked pattern of PNC. A pattern of PNC can also be caused indirectly, as a by-product of other causes, such as extinction, dispersal limitation, competition and predation. Recognition of patterns of PNC can contribute to understanding macroevolutionary processes: for example, release from constraint in traits has been hypothesized to trigger adaptive radiations such as that of the angiosperms. Given the multiple causes of patterns of PNC, tests should address explicit questions about hypothesized processes. We conclude that PNC is a scientifically useful concept with applications to the practice of ecological research.

Although arbuscular mycorrhizal (AM) fungi are obligate symbionts that can influence plant growth, the magnitude and direction of these effects are highly variable within fungal genera and even among isolates within species, as well as among plant taxa. To determine whether variability in AM fungal morphology and growth is correlated with AM fungal effects on plant growth, we established a common garden experiment with 56 AM fungal isolates comprising 17 genera and six families growing with three plant host species. Arbuscular mycorrhizal fungal morphology and growth was highly conserved among isolates of the same species and among species within a family. By contrast, plant growth response to fungal inoculation was highly variable, with the majority of variation occurring among different isolates of the same AM fungal species. Our findings show that host performance cannot be predicted from AM fungal morphology and growth traits. Divergent effects on plant growth among isolates within an AM fungal species may be caused by coevolution between co-occurring fungal and plant populations.

Taxa co‐occurring in communities often represent a nonrandom sample, in phenotypic or phylogenetic terms, of the regional species pool. While heuristic arguments have identified processes that create community phylogenetic patterns, further progress hinges on a more comprehensive understanding of the interactions between underlying ecological and evolutionary processes. We created a simulation framework to model trait evolution, assemble communities (via competition, habitat filtering, or neutral assembly), and test the phylogenetic pattern of the resulting communities. We found that phylogenetic community structure is greatest when traits are highly conserved and when multiple traits influence species membership in communities. Habitat filtering produces stronger phylogenetic structure when taxa with derived (as opposed to ancestral) traits are favored in the community. Nearest‐relative tests have greater power to detect patterns due to competition, while total community relatedness tests perform better with habitat filtering. The size of the local community relative to the regional pool strongly influences statistical power; in general, power increases with larger pool sizes for communities created by filtering but decreases for communities created by competition. Our results deepen our understanding of processes that contribute to phylogenetic community structure and provide guidance for the design and interpretation of empirical research.

Species distributions are often constrained by climatic tolerances that are ultimately determined by evolutionary history and/or adaptive capacity, but these factors have rarely been partitioned. Here, we experimentally determined two key climatic niche traits (desiccation and cold resistance) for 92—95 Drosophila species and assessed their importance for geographic distributions, while controlling for acclimation, phylogeny, and spatial autocorrelation. Employing an array of phylogenetic analyses, we documented moderate-to-strong phylogenetic signal in both desiccation and cold resistance. Desiccation and cold resistance were clearly linked to species distributions because significant associations between traits and climatic variables persisted even after controlling for phylogeny. We used different methods to untangle whether phylogenetic signal reflected phylogenetically related species adapted to similar environments or alternatively phylogenetic inertia. For desiccation resistance, weak phylogenetic inertia was detected; ancestral trait reconstruction, however, revealed a deep divergence that could be traced back to the genus level. Despite drosophilids' high evolutionary potential related to short generation times and high population sizes, cold resistance was found to have a moderate-to-high level of phylogenetic inertia, suggesting that evolutionary responses are likely to be slow. Together these findings suggest species distributions are governed by evolutionarily conservative climate responses, with limited scope for rapid adaptive responses to future climate change.

Traits do not evolve in isolation but often as part of integrated trait syndromes, yet the relative contributions of environmental effects and evolutionary history on traits and their correlations are not easily resolved. In the present study, we develop a methodological framework to elucidate eco‐evolutionary patterns in functional trait syndromes. We do so by separating the amount of variance and covariance related to phylogenetic heritage and environmental variables ( environmental phylogenetic conservatism ), only phylogenetic heritage ( non‐attributed phylogenetic conservatism ) and only to environmental variables ( evolutionarily labile environmental effects ). Variance–covariance structures of trait syndromes are displayed as networks. We then use this framework to guide a newly derived imputation method based on machine learning models that predict trait values for unsampled taxa, considering environmental and phylogenetic information as well as trait covariation. TrEvol is presented as an R package providing a unified set of methodologies to study and predict multivariate trait patterns and improve our capacity to impute trait values. To illustrate its use, we leverage both simulated data and species‐level traits related to hydraulics and the leaf economics spectrum, in relation to an aridity index, demonstrating that most trait correlations can be attributed to environmental phylogenetic conservatism . This conceptual framework can be employed to examine issues ranging from the evolution of trait adaptation at different phylogenetic depths to intraspecific trait variation.