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Abstract Gymnosperms encompass a diverse group of mostly woody plants with high ecological and economic value, yet little is known about the scope and organization of fine‐root trait diversity among gymnosperms due to the undersampling of most gymnosperm families and the dominance of angiosperm groups in recent syntheses.New and existing data were compiled for morphological traits (root diameter, length, tissue density, specific root length [SRL] and specific root area [SRA]), the architectural trait branching ratio, root nitrogen content [N] and mycorrhizal colonization. We used phylogenetic least squares regression and principal component analysis to determine trait–trait relationships and coordination across 66 species, representing 11 of the 12 extant gymnosperm families from boreal, temperate, subtropical and tropical biomes. Finally, we compared the relationship between family divergence time and mean trait values to determine whether evolutionary history structured variation in fine‐root traits within the gymnosperm phylogeny.Wide variation in gymnosperm root traits could be largely captured by two primary axes of variation defined by SRL and diameter, and root tissue density and root nitrogen, respectively. However, individual root length and SRA each had significant correlations with traits defining both main axes of variation. Neither mycorrhizal colonization nor root branching ratio were closely related to other traits. We did not observe a directional evolution of mean trait values from older to more recently diverged gymnosperm families.Synthesis. Despite their unique evolutionary history, gymnosperms display a root economic space similar to that identified in angiosperms, likely reflecting common constraints on plants adapting to diverse environments in both groups. These findings provide greater confidence that patterns observed in broad syntheses justly capture patterns of trait diversity among multiple, distinct lineages. Additionally, independence between root architecture and other traits may support greater diversity in below‐ground resource acquisition strategies. Unlike angiosperms, there were no clear trends towards increasingly thin roots over evolutionary time, possibly because of lower diversification rates or unique biogeographic history among gymnosperms, though additional observations are needed to more richly test evolutionary trends among gymnosperms.
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Identifying direct and indirect associations among traits by merging phylogenetic comparative methods and structural equation models
Abstract Traits underlie organismal responses to their environment and are essential to predict community responses to environmental conditions under global change. Species differ in life‐history traits, morphometrics, diet type, reproductive characteristics and habitat utilization.Trait associations are widely analysed using phylogenetic comparative methods (PCM) to account for correlations among related species. Similarly, traits are measured for some but not all species, and missing continuous traits (e.g. growth rate) can be imputed using ‘phylogenetic trait imputation’ (PTI), based on evolutionary relatedness and trait covariance. However, PTI has not been available for categorical traits, and estimating covariance among traits without ecological constraints risks inferring implausible evolutionary mechanisms.Here, we extend previous PCM and PTI methods by (1) specifying covariance among traits as a structural equation model (SEM), and (2) incorporating associations among both continuous and categorical traits. Fitting a SEM replaces the covariance among traits with a set of linear path coefficients specifying potential evolutionary mechanisms. Estimated parameters then represent regression slopes (i.e. the average change in trait Y given an exogenous change in trait X) that can be used to calculate both direct effects (X impacts Y) and indirect effects (X impacts Z and Z impacts Y).We demonstrate phylogenetic structural‐equation mixed‐trait imputation using 33 variables representing life history, reproductive, morphological, and behavioural traits for all >32,000 described fishes worldwide. SEM coefficients suggest that one degree Celsius increase in habitat is associated with an average 3.5% increase in natural mortality (including a 1.4% indirect impact that acts via temperature effects on the growth coefficient), and an average 3.0% decrease in fecundity (via indirect impacts on maximum age and length). Cross‐validation indicates that the model explains 54%–89% of variance for withheld measurements of continuous traits and has an area under the receiver‐operator‐characteristics curve of 0.86–0.99 for categorical traits.We use imputed traits to classify all fishes into life‐history types, and confirm a phylogenetic signal in three dominant life‐history strategies in fishes. PTI using phylogenetic SEMs ensures that estimated parameters are interpretable as regression slopes, such that the inferred evolutionary relationships can be compared with long‐term evolutionary and rearing experiments.
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- Award ID(s):
- 2109411
- PAR ID:
- 10403270
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Methods in Ecology and Evolution
- Volume:
- 14
- Issue:
- 5
- ISSN:
- 2041-210X
- Page Range / eLocation ID:
- p. 1259-1275
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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