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Deciphering the global distribution of polyploid plants is fundamental for understanding plant evolution and ecology. Many factors have been hypothesized to affect the uneven distribution of polyploid plants across the globe. Nevertheless, the lack of large comparative datasets has restricted such studies to local floras and to narrow taxonomical scopes, limiting our understanding of the underlying drivers of polyploid plant distribution. We present a map portraying the worldwide polyploid frequencies, based on extensive spatial data coupled with phylogeny-based polyploidy inference for tens of thousands of species. This allowed us to assess the potential global drivers affecting polyploid distribution. Our data reveal a clear latitudinal trend, with polyploid frequency increasing away from the equator. Climate, especially temperature, appears to be the most influential predictor of polyploid distribution. However, we find this effect to be mostly indirect, mediated predominantly by variation in plant lifeforms and, to a lesser extent, by taxonomical composition and species richness. Thus, our study presents an emerging view of polyploid distribution that highlights attributes that facilitate the establishment of new polyploid lineages by providing polyploids with sufficient time (that is, perenniality) and space (low species richness) to compete with pre-adapted diploid relatives.

Hybridization and genome doubling (allopolyploidy) have led to evolutionary novelties as well as to the origin of new clades and species. Despite the importance of allopolyploidization, the dynamics of postpolyploid diploidization (PPD) at the genome level has been only sparsely studied. The Microlepidieae (MICR) is a crucifer tribe of 17 genera and c. 56 species endemic to Australia and New Zealand. Our phylogenetic and cytogenomic analyses revealed that MICR originated via an intertribal hybridization between ancestors of Crucihimalayeae (n = 8; maternal genome) and Smelowskieae (n = 7; paternal genome), both native to the Northern Hemisphere. The reconstructed ancestral allopolyploid genome (n = 15) originated probably in northeastern Asia or western North America during the Late Miocene (c. 10.6-7 million years ago) and reached the Australian mainland via long-distance dispersal. In Australia, the allotetraploid genome diverged into at least three main subclades exhibiting different levels of PPD and diversity: 1.25-fold descending dysploidy (DD) of n = 15 → n = 12 (autopolyploidy → 24) in perennial Arabidella (3 species), 1.5-fold DD of n = 15 → n = 10 in the perennial Pachycladon (11 spp.) and 2.1-3.75-fold DD of n = 15 → n = 7-4 in the largely annual crown-group genera (42 spp. in 15 genera). These results are among the first to demonstrate multispeed genome evolution in taxa descending from a common allopolyploid ancestor. It is suggested that clade-specific PPD can operate at different rates and efficacies and can be tentatively linked to life histories and the extent of taxonomic diversity., (© 2017 John Wiley & Sons Ltd.)

Premise of the Study: Flowering plants display a variety of sexual systems, ranging from complete cosexuality (hermaphroditism) to separate-sexed individuals (dioecy). While dioecy is relatively rare, it has evolved many times and is present in many plant families. Transitions in sexual systems are hypothesized to be affected by large genomic events such as whole-genome duplication, or polyploidy, and several models have been proposed to explain the observed patterns of association., Methods: In this study, we assessed the association between ploidy and sexual system (separate or combined sexes). To this end, we assembled a database of ploidy levels and sexual systems for ∼1000 species, spanning 18 genera and 15 families. We applied several phylogenetic comparative approaches, including Pagel's coevolutionary framework and sister clade analyses, for detecting correlations between ploidy level and sexual system., Key Results: Our results indicate a broad association between polyploidy and sexual system dimorphism, with low evolutionary stability of the diploid-dioecious condition observed in several clades. A detailed examination of the clades exhibiting this correlation reveals that it is underlain by various patterns of transition rate asymmetry., Conclusions: We conclude that the long-hypothesized connection between ploidy and sexual system holds in some clades, although it may well be affected by factors that differ from clade to clade. Our results further demonstrate that to better understand the evolutionary processes involved, more sophisticated methods and extensive and detailed data sets are required for both broad and focused inquiry., (© 2016 Botanical Society of America.)

Premise of the Study: Polyploidization is a common and recurring phenomenon in plants and is often thought to be a mechanism of "instant speciation". Whether polyploidization is associated with the formation of new species (cladogenesis) or simply occurs over time within a lineage (anagenesis), however, has never been assessed systematically., Methods: We tested this hypothesis using phylogenetic and karyotypic information from 235 plant genera (mostly angiosperms). We first constructed a large database of combined sequence and chromosome number data sets using an automated procedure. We then applied likelihood models (ClaSSE) that estimate the degree of synchronization between polyploidization and speciation events in maximum likelihood and Bayesian frameworks., Key Results: Our maximum likelihood analysis indicated that 35 genera supported a model that includes cladogenetic transitions over a model with only anagenetic transitions, whereas three genera supported a model that incorporates anagenetic transitions over one with only cladogenetic transitions. Furthermore, the Bayesian analysis supported a preponderance of cladogenetic change in four genera but did not support a preponderance of anagenetic change in any genus., Conclusions: Overall, these phylogenetic analyses provide the first broad confirmation that polyploidization is temporally associated with speciation events, suggesting that it is indeed a major speciation mechanism in plants, at least in some genera., (© 2016 Botanical Society of America.)

Polyploidy, the doubling of genomic content, is a widespread feature, especially among plants, yet its macroevolutionary impacts are contentious. Traditionally, polyploidy has been considered an evolutionary dead end, whereas recent genomic studies suggest that polyploidy has been a key driver of macroevolutionary success. We examined the consequences of polyploidy on the time scale of genera across a diverse set of vascular plants, encompassing hundreds of inferred polyploidization events. Likelihood-based analyses indicate that polyploids generally exhibit lower speciation rates and higher extinction rates than diploids, providing the first quantitative corroboration of the dead-end hypothesis. The increased speciation rates of diploids can, in part, be ascribed to their capacity to speciate via polyploidy. Only particularly fit lineages of polyploids may persist to enjoy longer-term evolutionary success.

Polyploidy, the genome wide duplication of chromosome number, is a key feature in eukaryote evolution. Polyploidy exists in diverse groups including animals, fungi, and invertebrates but is especially prevalent in plants with most, if not all, plant species having descended from a polyploidization event. Polyploids often differ markedly from their diploid progenitors in morphological, physiological, and life history characteristics as well as rates of adaptation. The altered characteristics displayed by polyploids may contribute to their success in novel ecological habitats. Clearly, a better understanding of the processes underlying changes in the number of chromosomes within genomes is a key goal in our understanding of speciation and adaptation for a wide range of families and genera. Despite the fundamental role of chromosome number change in eukaryotic evolution, probabilistic models describing the evolution of chromosome number along a phylogeny have not yet been formulated. We present a series of likelihood models, each representing a different hypothesis regarding the evolution of chromosome number along a given phylogeny. These models allow us to reconstruct ancestral chromosome numbers and to estimate the expected number of polyploidization events and single chromosome changes (dysploidy) that occurred along a phylogeny. We test, using simulations, the accuracy of this approach and its dependence on the number of taxa and tree length. We then demonstrate the application of the method for the study of chromosome number evolution in 4 plant genera: Aristolochia, Carex, Passiflora, and Helianthus. Considering the depth of the available cytological and phylogenetic data, formal models of chromosome number evolution are expected to advance significantly our understanding of the importance of polyploidy and dysploidy across different taxonomic groups.

Since its discovery in 1907, polyploidy has been recognized as an important phenomenon in vascular plants, and several lines of evidence indicate that most, if not all, plant species ultimately have a polyploid ancestry. However, previous estimates of the frequency of polyploid speciation suggest that the formation and establishment of neopolyploid species is rare. By combining information from the botanical community's vast cytogenetic and phylogenetic databases, we establish that 15% of angiosperm and 31% of fern speciation events are accompanied by ploidy increase. These frequency estimates are higher by a factor of four than earlier estimates and lead to a standing incidence of polyploid species within genera of 35% (n = 1,506). Despite this high incidence, we find no direct evidence that polyploid lines, once established, enjoy greater net species diversification. Thus, the widespread occurrence of polyploid taxa appears to result from the substantial contribution of polyploidy to cladogenesis, but not from subsequent increases in diversification rates of polyploid lines.

Premise of research. Polyploidy, a major evolutionary process in flowering plants, is expected to impact floral traits, which can have cascading effects on pollination interactions, but this may depend on selfing propensity. In a novel use of herbarium specimens, we assessed the effects of polyploidy and mating system on floral traits and the pollination niche of 40 Brassicaceae species. Methodology. We combined data on mating system (self-compatible or self-incompatible) with inferred ploidy level (polyploid or diploid) and used phylogenetically controlled analyses to investigate their influence on floral traits (size and shape) and the degree of pollination generalism based on the frequency and the richness of heterospecific pollen morphospecies captured by stigmas. Pivotal results. Flower size (but not shape) depended on the interaction between ploidy and mating system. Self-incompatible polyploid species had larger flowers than self-incompatible diploids, but there was no difference for self-compatible species. The breadth of pollination niche (degree of generalism) was not affected by ploidy but was rather strongly affected by mating system only. Self-incompatible species had more stigmas with heterospecific pollen and higher heterospecific pollen morphospecies richness per stigma than self-compatible species, regardless of their ploidy. Conclusions. Our results demonstrate that mating system moderated the influence of ploidy on morphological features associated with pollination generalism but that response in terms of heterospecific pollen captured as a proxy of pollination generalism was more variable. [ABSTRACT FROM AUTHOR]

See also the Commentary on this article by Spoelhof et al., 240: 909–911. [ABSTRACT FROM AUTHOR]

Summary: Changes in chromosome numbers, including polyploidy and dysploidy events, play a key role in eukaryote evolution as they could expediate reproductive isolation and have the potential to foster phenotypic diversification. Deciphering the pattern of chromosome‐number change within a phylogeny currently relies on probabilistic evolutionary models. All currently available models assume time homogeneity, such that the transition rates are identical throughout the phylogeny.Here, we develop heterogeneous models of chromosome‐number evolution that allow multiple transition regimes to operate in distinct parts of the phylogeny. The partition of the phylogeny to distinct transition regimes may be specified by the researcher or, alternatively, identified using a sequential testing approach. Once the number and locations of shifts in the transition pattern are determined, a second search phase identifies regimes with similar transition dynamics, which could indicate on convergent evolution.Using simulations, we study the performance of the developed model to detect shifts in patterns of chromosome‐number evolution and demonstrate its applicability by analyzing the evolution of chromosome numbers within the Cyperaceae plant family.The developed model extends the capabilities of probabilistic models of chromosome‐number evolution and should be particularly helpful for the analyses of large phylogenies that include multiple distinct subclades. [ABSTRACT FROM AUTHOR]

Summary: Chromosome number is a central feature of eukaryote genomes. Deciphering patterns of chromosome‐number change along a phylogeny is central to the inference of whole genome duplications and ancestral chromosome numbers. ChromEvol is a probabilistic inference tool that allows the evaluation of several models of chromosome‐number evolution and their fit to the data. However, fitting a model does not necessarily mean that the model describes the empirical data adequately. This vulnerability may lead to incorrect conclusions when model assumptions are not met by real data.Here, we present a model adequacy test for likelihood models of chromosome‐number evolution. The procedure allows us to determine whether the model can generate data with similar characteristics as those found in the observed ones.We demonstrate that using inadequate models can lead to inflated errors in several inference tasks. Applying the developed method to 200 angiosperm genera, we find that in many of these, the best‐fitting model provides poor fit to the data. The inadequacy rate increases in large clades or in those in which hybridizations are present.The developed model adequacy test can help researchers to identify phylogenies whose underlying evolutionary patterns deviate substantially from current modelling assumptions and should guide future methods development. [ABSTRACT FROM AUTHOR]

Chromosome numbers have been widely used to describe the most fundamental genomic attribute of an organism or a lineage. Although providing strong phylogenetic signal, chromosome numbers vary remarkably among eukaryotes at all levels of taxonomic resolution. Changes in chromosome numbers regularly serve as indication of major genomic events, most notably polyploidy and dysploidy. Here, we review recent advancements in our ability to make inferences regarding historical events that led to alterations in the number of chromosomes of a lineage. We first describe the mechanistic processes underlying changes in chromosome numbers, focusing on structural chromosomal rearrangements. Then, we focus on experimental procedures, encompassing comparative cytogenomics and genomics approaches, and on computational methodologies that are based on explicit models of chromosome-number evolution. Together, these tools offer valuable predictions regarding historical events that have changed chromosome numbers and genome structures, as well as their phylogenetic and temporal placements. [ABSTRACT FROM AUTHOR]

Local adaptation of plants along environmental gradients provides strong evidence for clinal evolution mediated by natural selection. Plants have developed diverse strategies to mitigate stress, for example, drought escape is a phenological strategy to avoid drought stress, while polyploidy was proposed as a genomic adaptation to stress. Polyploidy as an adaptation to aridity (an environmental parameter integrating temperature and precipitation) was previously documented in annual Brachypodium spp. (Poaceae) in the Western Mediterranean. Here, we examined whether polyploidy or phenology are associated with aridity in annual Brachypodium spp. along the aridity gradient in the Eastern Mediterranean. Using flow cytometry, we determined ploidy levels of plants from natural populations along the Israeli gradient, spanning ∼424 km from mesic Mediterranean to extreme desert climates. In a common garden we recorded time of seedling emergence, flowering and senescence. We tested whether the proportion of allotetraploids in the populations and phenological traits were associated with aridity. Contrary to a previous study in the Western Mediterranean, we found no effect of aridity on the proportion of allotetraploids and diploids within populations. Interestingly, phenology was associated with aridity: time of emergence was later, while flowering and senescence were earlier in desert plants. Our results indicate that in the Eastern Mediterranean, adaptation of Brachypodium to aridity is mediated mainly by phenology, rather than ploidy level. Therefore, we suggest that genome duplication is not the main driver of adaptation to environmental stress; rather, phenological change as a drought escape mechanism may be the major adaptation. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Species diversity is remarkably unevenly distributed among flowering plant lineages. Despite a growing toolbox of research methods, the reasons underlying this patchy pattern have continued to perplex plant biologists for the past two decades. In this review, we examine the present understanding of transitions in flowering plant evolution that have been proposed to influence speciation and extinction. In particular, ploidy changes, transitions between tropical and nontropical biomes, and shifts in floral form have received attention and have offered some surprises in terms of which factors influence speciation and extinction rates. Mating systems and dispersal characteristics once predominated as determining factors, yet recent evidence suggests that these changes are not as influential as previously thought or are important only when paired with range shifts. Although range extent is an important correlate of speciation, it also influences extinction and brings an applied focus to diversification research. Recent studies that find that past diversification can predict present-day extinction risk open an exciting avenue for future research to help guide conservation prioritization. [ABSTRACT FROM AUTHOR]

The author discusses the Chromosome Counts Database (CCDB), which is an information resource about plant chromosome numbers. Topics discussed include the evolutionary pattern of chromosome number, the use of chromosome numbers as an important phylogenetic character in cytotaxonomy, and the Index to Plant Chromosome Numbers (IPCN).