Summary The effects of single chromosome number change—dysploidy – mediating diversification remain poorly understood. Dysploidy modifies recombination rates, linkage, or reproductive isolation, especially for one‐fifth of all eukaryote lineages with holocentric chromosomes. Dysploidy effects on diversification have not been estimated because modeling chromosome numbers linked to diversification with heterogeneity along phylogenies is quantitatively challenging.We propose a new state‐dependent diversification model of chromosome evolution that links diversification rates to dysploidy rates considering heterogeneity and differentiates between anagenetic and cladogenetic changes. We apply this model toCarex(Cyperaceae), a cosmopolitan flowering plant clade with holocentric chromosomes.We recover two distinct modes of chromosomal evolution and speciation inCarex. In one diversification mode, dysploidy occurs frequently and drives faster diversification rates. In the other mode, dysploidy is rare, and diversification is driven by hidden, unmeasured factors. When we use a model that excludes hidden states, we mistakenly infer a strong, uniformly positive effect of dysploidy on diversification, showing that standard models may lead to confident but incorrect conclusions about diversification.This study demonstrates that dysploidy can have a significant role in speciation in a large plant clade despite the presence of other unmeasured factors that simultaneously affect diversification.
more »
« less
Simpson's tachytely or bradytely? The importance of quantifying rate uncertainty
Abstract The spectacular variation in species forms and richness across space and time can be explored using sophisticated and powerful tools recently developed by evolutionary modellers. In this contribution, we ask if the classic ‘Simpsonian’ view of tachytelic (fast), horotelic (standard) and bradytelic (slow) diversification rates can be distinguished with currently available tools and data. A neglected topic here is the role that the uncertainty of diversification rate estimates plays, where the lack of in‐depth uncertainty measures could hinder our ability to confidently suggest differences in speciation or extinction rates in any given comparison.We propose quantifying the relative uncertainty of diversification estimates, to better compare diversification tempo across phylogenies of different sizes and ages. We present three case studies, using the most popular models for diversification rate estimation, with or without fossils, to investigate claims of bradytely or tachytely. Using summary statistics and linear models, we ask if point estimates of diversification rates are comparable across clades. More specifically, we fit a linear model to understand which phylogenetic tree properties (including size and age) may affect the uncertainty of diversification estimates.We found the ‘Goldilocks of uncertainty’: Phylogenies that are young with insufficient tips or that are old increase the uncertainty of diversification estimates. The choice of diversification modelling approach is independent of the pattern of diversification rates decaying exponentially with clade age.In practice, we still cannot confidently compare diversification rates or their variation, due to uncertainties stemming from clade age, sample size and biased sampling. We emphasize the need for researchers to focus on estimating and presenting uncertainty in their estimates. Such uncertainty estimates are currently absent from many publications, limiting our ability to compare the tempo of diversifications across the tree of life. We conclude by proposing solutions and guidelines to encourage new studies for measure uncertainty.
more »
« less
- Award ID(s):
- 2323170
- PAR ID:
- 10556185
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Methods in Ecology and Evolution
- Volume:
- 16
- Issue:
- 1
- ISSN:
- 2041-210X
- Format(s):
- Medium: X Size: p. 4-18
- Size(s):
- p. 4-18
- Sponsoring Org:
- National Science Foundation
More Like this
-
-
Summary Shifts in the age or turnover time of non‐structural carbohydrates (NSC) may underlie changes in tree growth under long‐term increases in drought stress associated with climate change. But NSC responses to drought are challenging to quantify, due in part to large NSC stores in trees and subsequently long response times of NSC to climate variation.We measured NSC age (Δ14C) along with a suite of ecophysiological metrics inPinus edulistrees experiencing either extreme short‐term drought (−90% ambient precipitation plot, 2020–2021) or a decade of severe drought (−45% plot, 2010–2021). We tested the hypothesis that carbon starvation – consumption exceeding synthesis and storage – increases the age of sapwood NSC.One year of extreme drought had no impact on NSC pool size or age, despite significant reductions in predawn water potential, photosynthetic rates/capacity, and twig and needle growth. By contrast, long‐term drought halved the age of the sapwood NSC pool, coupled with reductions in sapwood starch concentrations (−75%), basal area increment (−39%), and bole respiration rates (−28%).Our results suggest carbon starvation takes time, as tree carbon reserves appear resilient to extreme disturbance in the short term. However, after a decade of drought, trees apparently consumed old stored NSC to support metabolism.more » « less
-
Abstract Evolutionary biologists characterize macroevolutionary trends of phenotypic change across the tree of life using phylogenetic comparative methods. However, within‐species variation can complicate such investigations. For this reason, procedures for incorporating nonstructured (random) intraspecific variation have been developed.Likewise, evolutionary biologists seek to understand microevolutionary patterns of phenotypic variation within species, such as sex‐specific differences or allometric trends. Additionally, there is a desire to compare such within‐species patterns across taxa, but current analytical approaches cannot be used to interrogate within‐species patterns while simultaneously accounting for phylogenetic non‐independence. Consequently, deciphering how intraspecific trends evolve remains a challenge.Here we introduce an extended phylogenetic generalized least squares (E‐PGLS) procedure which facilitates comparisons of within‐species patterns across species while simultaneously accounting for phylogenetic non‐independence.Our method uses an expanded phylogenetic covariance matrix, a hierarchical linear model, and permutation methods to obtain empirical sampling distributions and effect sizes for model effects that can evaluate differences in intraspecific trends across species for both univariate and multivariate data, while conditioning them on the phylogeny.The method has appropriate statistical properties for both balanced and imbalanced data. Additionally, the procedure obtains evolutionary covariance estimates that reflect those from existing approaches for nonstructured intraspecific variation. Importantly, E‐PGLS can detect differences in structured (i.e. microevolutionary) intraspecific patterns across species when such trends are present. Thus, E‐PGLS extends the reach of phylogenetic comparative methods into the intraspecific comparative realm, by providing the ability to compare within‐species trends across species while simultaneously accounting for shared evolutionary history.more » « less
-
Summary If particular traits consistently affect rates of speciation and extinction, broad macroevolutionary patterns can be interpreted as consequences of selection at high levels of the biological hierarchy. Identifying traits associated with diversification rates is difficult because of the wide variety of characters under consideration and the statistical challenges of testing for associations from comparative phylogenetic data. Ploidy (diploid vs polyploid states) and breeding system (self‐incompatible vs self‐compatible states) are both thought to be drivers of differential diversification in angiosperms.We fit 29 diversification models to extensive trait and phylogenetic data in Solanaceae and investigate how speciation and extinction rate differences are associated with ploidy, breeding system, and the interaction between these traits.We show that diversification patterns in Solanaceae are better explained by breeding system and an additional unobserved factor, rather than by ploidy. We also find that the most common evolutionary pathway to polyploidy in Solanaceae occurs via direct breakdown of self‐incompatibility by whole genome duplication, rather than indirectly via breakdown followed by polyploidization.Comparing multiple stochastic diversification models that include complex trait interactions alongside hidden states enhances our understanding of the macroevolutionary patterns in plant phylogenies.more » « less
-
Abstract Understanding the mechanisms that promote the coexistence of hundreds of species over small areas in tropical forest remains a challenge. Many tropical tree species are presumed to be functionally equivalent shade tolerant species but exist on a continuum of performance trade‐offs between survival in shade and the ability to quickly grow in sunlight. These trade‐offs can promote coexistence by reducing fitness differences.Variation in plant functional traits related to resource acquisition is thought to predict variation in performance among species, perhaps explaining community assembly across habitats with gradients in resource availability. Many studies have found low predictive power, however, when linking trait measurements to species demographic rates.Seedlings face different challenges recruiting on the forest floor and may exhibit different traits and/or performance trade‐offs than older individuals face in the eventual adult niche. Seed mass is the typical proxy for seedling success, but species also differ in cotyledon strategy (reserve vs. photosynthetic) or other leaf, stem and root traits. These can cause species with the same average seed mass to have divergent performance in the same habitat.We combined long‐term studies of seedling dynamics with functional trait data collected at a standard life‐history stage in three diverse neotropical forests to ask whether variation in coordinated suites of traits predicts variation among species in demographic performance.Across hundreds of species in Ecuador, Panama and Puerto Rico, we found seedlings displayed correlated suites of leaf, stem, and root traits, which strongly correlated with seed mass and cotyledon strategy. Variation among species in seedling functional traits, seed mass, and cotyledon strategy were strong predictors of trade‐offs in seedling growth and survival. These results underscore the importance of matching the ontogenetic stage of the trait measurement to the stage of demographic dynamics.Our findings highlight the importance of cotyledon strategy in addition to seed mass as a key component of seed and seedling biology in tropical forests because of the contribution of carbon reserves in storage cotyledons to reducing mortality rates and explaining the growth‐survival trade‐off among species.Synthesis: With strikingly consistent patterns across three tropical forests, we find strong evidence for the promise of functional traits to provide mechanistic links between seedling form and demographic performance.more » « less
An official website of the United States government
