Diversification Rates Research Articles

The evolution of chromosome number during the diversification of the tribe Vernonieae (Asteraceae)

Abstract Changes in chromosome number have played an important role in the diversification and evolution of angiosperms. In Asteraceae, tribe Vernonieae are one of the most variable groups with regard to chromosome number. Previously, chromosome numbers n = 9 and 10 were thought to characterize the Old World members of the tribe, and n = 14, 16, 17 and 18 the New World members. This scenario was revised as a result of reports of new chromosome numbers, but the events leading to this wide variation remain unknown. Here we carried out a phylogenetic analysis of Vernonieae in a temporal framework, assessing patterns of diversification and establishing possible relationships with chromosome events. Chromosomal evolution was analysed with ChromEvol, from a phylogenetic tree dated in BEAST. Shifts in diversification rates using Bayesian analysis of macroevolutionary mixtures were inferred. Vernonieae originated ~46 Mya and the diversification rate increased sharply ~11 Mya after the Mid-Miocene Climatic Optimum. The ancestral chromosome number for the tribe was n = 10, which remained stable for Old World taxa, whereas n = 9 was the ancestral number for New World species. The tribe has undergone 32 chromosome rearrangements throughout its evolutionary history, with dysploidy and polyploidy possibly explaining the observed diversification pattern.

Why are there so many species of mining bees (Hymenoptera, Andrenidae)? The possible roles of phenology and Wolbachia incompatibility in maintaining species boundaries in the Andrena proxima‐complex

AbstractThe factors leading to the extraordinary diversity of the bees of the genus Andrena Fabricius, the second most speciose genus among bees, remain largely unknown. To examine the pattern of diversification in this genus, we investigate species boundaries within the controversial Andrena proxima‐complex using a dataset of ultra‐conserved elements and various species delimitation analyses (admixture analyses, BPP, DAPC). Our results confirm the presence of three separate species in this species group (Andrena proxima (Kirby), A. ampla Warncke and A. alutacea Stöckhert) and suggest very low levels of interspecific gene flow. Andrena proxima and A. alutacea are regularly found in sympatry, suggesting an advanced stage of speciation. By contrast, A. ampla shows a parapatric distribution with both other species. Andrena alutacea harbours a unique strain of Wolbachia Hertig, absent in the other two species, which are infected by the two same strains of Wolbachia. In addition, the three species have distinct phenologies, with A. proxima flying earlier in the season and A. alutacea significantly later; the phenology of A. ampla is intermediate. Our results suggest that Wolbachia‐induced incompatibilities and phenological differences maintain species boundaries in this group. The most advanced stage of speciation is observed in the two species showing putatively incompatible strains of Wolbachia and strongly distinct phenologies, A. alutacea and A. proxima. We propose that the smaller differences in phenological differentiation between A. ampla and both other species may explain the observed pattern of parapatric distribution. We discuss how these factors may underlie the high diversification rate observed in other groups of Andrena, a genus characterized by particularly variable and short flying periods.

Flexible drought deciduousness in a neotropical understory herb.

PremiseAdaptive divergence across environmental gradients is a key driver of speciation. Precipitation seasonality gradients are common in the tropics, yet drought adaptation is nearly unexplored in neotropical understory herbs. Here, we examined two recently diverged neotropical spiral gingers, one adapted to seasonal drought and one reliant on perennial water, to uncover the basis of drought adaptation.MethodsWe combined ecophysiological trait measurements in the field and greenhouse with experimental and observational assessments of real‐time drought response to determine how Costus villosissimus (Costaceae) differs from C. allenii to achieve drought adaptation.ResultsWe found that drought‐adapted C. villosissimus has several characteristics indicating flexible dehydration avoidance via semi‐drought‐deciduousness and a fast economic strategy. Although the two species do not differ in water‐use efficiency, C. villosissimus has a more rapid growth rate, lower leaf mass per area, lower stem density, higher leaf nitrogen, and a strong trend of greater light‐saturated photosynthetic rates. These fast economic strategy traits align with both field‐based observations and experimental dry‐down results. During drought, C. villosissimus displays facultative drought‐deciduousness, losing lower leaves during the dry season and rapidly growing new leaves in the wet season.ConclusionsWe revealed a drought adaptation strategy that has not, to our knowledge, previously been documented in tropical herbs. This divergent drought adaptation evolved recently and is an important component of reproductive isolation between C. villosissimus and C. allenii, indicating that adaptive shifts to survive seasonal drought may be an underappreciated axis of neotropical understory plant diversification.

Time for speciation and niche conservatism explain the latitudinal diversity gradient in clupeiform fishes

AbstractAimThe latitudinal diversity gradient of increasing species richness from poles to equator is one of the most striking and pervasive spatial patterns of biodiversity. Climate appears to have been key to the formation of the latitudinal diversity gradient, but the processes through which climate shaped species richness remain unclear. We tested predictions of the time for speciation, carrying capacity and diversification rate latitudinal diversity gradient hypotheses in a trans‐marine/freshwater clade of fishes.LocationGlobal in marine and freshwater environments.TaxonClupeiformes (anchovies, herrings, sardines and relatives).MethodsWe tested predictions of latitudinal diversity gradient hypotheses using a molecular phylogeny, species distribution data and phylogenetic comparative approaches. To test the time for speciation hypothesis, we conducted ancestral state reconstructions to infer the ages of temperate, subtropical and tropical lineages and frequency of evolutionary transitions between climates. We tested the carry capacity hypothesis by characterizing changes in net diversification rates through time. To test the diversification rate hypothesis, we qualitatively compared the diversification rates of temperate, subtropical and tropical lineages and conducted statistical tests for associations between latitude and diversification rates.ResultsWe identified four transitions to temperate climates and two transitions out of temperate climates. We found no differences in diversification rates among temperate and tropical clupeiforms. Net diversification rates remained positive in crown Clupeiformes since their origin ~150 Ma in both tropical and temperate lineages. Climate niche characters exhibited strong phylogenetic signal. All temperate clupeiform lineages arose <50 Ma, after the Early Eocene Climatic Optimum.Main conclusionsOur results support the time for speciation hypothesis, which proposes that climate niche conservatism and fluctuations in the extent of temperate climates limited the time for species to accumulate in temperate climates, resulting in the latitudinal diversity gradient. We found no support for the carrying capacity or diversification rate hypotheses.

Diversification rates in Tardigrada indicate a decreasing tempo of lineage splitting regardless of reproductive mode

Understanding the dynamics of speciation and extinction events is one of the most interesting subjects in evolutionary biology that relates to all life forms, even the smallest ones. Tardigrades are microscopic invertebrates that attracted public and scientific attention mostly due to their ability to enter into the diapause stage called cryptobiosis and in such stage resist extremely harsh environmental conditions. However, although recent research solved a considerable number of phylogenetic uncertainties and further uncovered physiological mechanisms of cryptobiosis, not much attention is given to the evolutionary forces shaping tardigrade diversity. Here, we investigated the effect of reproductive mode on diversification rates in tardigrades using three groups: macrobiotids, echiniscids and milnesids, which represent low, moderate and high levels of parthenogenesis, respectively. Our results showed a decreasing tempo of diversification events for each of the studied groups without any differences that could be ascribed to reproductive mode. We discussed the observed lack of effect in tardigrades acknowledging deficiencies in available data sets and encouraging further studies to understand whether our results can be considered reliable.

Biogeographic history of a large clade of ectomycorrhizal fungi, the Russulaceae, in the Neotropics and adjacent regions.

The biogeography of neotropical fungi remains poorly understood. Here, we reconstruct the origins and diversification of neotropical lineages in one of the largest clades of ectomycorrhizal fungi in the globally widespread family Russulaceae. We inferred a supertree of 3285 operational taxonomic units, representing worldwide internal transcribed spacer sequences. We reconstructed biogeographic history and diversification and identified lineages in the Neotropics and adjacent Patagonia. The ectomycorrhizal Russulaceae have a tropical African origin. The oldest lineages in tropical South America, most with African sister groups, date to the mid-Eocene, possibly coinciding with a boreotropical migration corridor. There were several transatlantic dispersal events from Africa more recently. Andean and Central American lineages mostly have north-temperate origins and are associated with North Andean uplift and the general north-south biotic interchange across the Panama isthmus, respectively. Patagonian lineages have Australasian affinities. Diversification rates in tropical South America and other tropical areas are lower than in temperate areas. Neotropical Russulaceae have multiple biogeographic origins since the mid-Eocene involving dispersal and co-migration. Discontinuous distributions of host plants may explain low diversification rates of tropical lowland ectomycorrhizal fungi. Deeply diverging neotropical fungal lineages need to be better documented.

Exploitation of an ancestral pheromone biosynthetic pathway contributes to diversification in Heliconius butterflies.

During courtship, male butterflies of many species produce androconial secretions containing male sex pheromones (MSPs) that communicate species identity and affect female choice. MSPs are thus likely candidates as reproductive barriers, yet their role in speciation remains poorly studied. Although Heliconius butterflies are a model system in speciation, their MSPs have not been investigated from a macroevolutionary perspective. We use GC/MS to characterize male androconial secretions in 33 of the 69 species in the Heliconiini tribe. We found these blends to be species-specific, consistent with a role in reproductive isolation. We detected a burst in blend diversification rate at the most speciose genus, Heliconius; a consequence of Heliconius and Eueides species using a fatty acid (FA) metabolic pathway to unlock more complex blends than basal Heliconiini species, whose secretions are dominated by plant-like metabolites. A comparison of 10 sister species pairs demonstrates a striking positive correlation between blend dissimilarity and range overlap, consistent with character displacement or reinforcement in sympatry. These results demonstrate for the first time that MSP diversification can promote reproductive isolation across this group of butterflies, showcasing how implementation of an ancestral trait, the co-option of the FA metabolic pathway for pheromone production, can facilitate rapid speciation.

Effects of Plasticity on Elevational Range Size and Species Richness.

AbstractElevational ranges within many taxa are greater in the north temperate region than the tropics. Two hypotheses to explain the pattern are, first, that large elevational ranges in the temperate region arise because species have evolved broad tolerance curves in response to seasonality and, second, that a low diversification rate in the temperate region (speciation minus extinction) has led to relatively few species, each of which occupies a large elevational range in the absence of competitors (character release). We build a quantitative genetic model of selection on a phenotypic trait, whereby increased tolerance is modeled as arising from plasticity in the trait. We show that broad tolerances result in evolution of large elevational ranges because they induce shallower genotypic clines and hence reduced maladaptive gene flow. The evolution of large elevational ranges results in relatively few competing species arranged along the elevational gradient at a species carrying capacity. In such saturated communities, species have much elevational overlap. In contrast, in similar-sized communities that could accommodate many more species, the resulting character release is associated with smaller elevational overlaps. Empirical assessment of these predictions should contribute to assessing any role for ecological limits in driving the latitudinal diversity gradient in species richness.

Rapid preservation of Jehol Biota in Northeast China from high precision 40Ar/39Ar geochronology

In Liaoning Province, Northeastern China, the Early Cretaceous Jehol Biota records significant evolution and radiation of fauna in terms of diversity, abundance and complexity. The alluvial-to-fluvial Lujiatun and lacustrine Jianshangou beds of the lower Yixian Formation yield exceptionally well-preserved three-dimensional and articulated dinosaur and mammal fossils, and two-dimensional primitive birds/feathered dinosaurs, respectively. Published 40Ar/39Ar ages for the two fossil beds have uncertainties of ± 0.4 to 1.0 Ma (± 2σ analytical uncertainty), and these reflect the measurement of large, and potentially heterogeneous, populations of feldspar crystals. More precise constraints on the timing and duration of biological radiation recorded by this sequence can further quantify evolutionary patterns and rates of diversification of important Early Cretaceous animals and plants. Single crystal 40Ar/39Ar fusion experiments on sanidine from intercalated ash beds yield dates that constrain the deposition of the Jianshangou and Lujiatun fossil beds to between 125.374 ± 0.085 and 125.572 ± 0.120 Ma, and 125.541 ± 0.099 and 125.793 ± 0.127 Ma, respectively. These ages indicate that the lower part of Jianshangou fossil bed was deposited nearly contemporaneously with the Lujiatun fossil bed. Thus, mass mortality events reflected by dinosaur fossil concentrations in the Lujiatun fossil bed and feathered dinosaur/primitive bird fossils in the Jianshangou fossil bed were nearly coeval and short-lived. These 40Ar/39Ar ages, combined with astrochronologic and paleomagnetic results from the lacustrine sediments in the Jianshangou fossil bed, not only refine the duration and rapid radiation of primitive birds and feathered dinosaurs in the deposit, but also suggest that the distribution of magnetic polarity chron boundaries within the Barremian stage in the current geologic time scale (GTS 2020) requires a revision.

A phylogenetic study to assess the link between biome specialization and diversification in swallowtail butterflies.

The resource‐use hypothesis, proposed by E.S. Vrba, states that habitat fragmentation caused by climatic oscillations would affect particularly biome specialists (species inhabiting only one biome), which might show higher speciation and extinction rates than biome generalists. If true, lineages would accumulate biome‐specialist species. This effect would be particularly exacerbated for biomes located at the periphery of the global climatic conditions, namely, biomes that have high/low precipitation and high/low temperature such as rainforest (warm‐humid), desert (warm‐dry), steppe (cold‐dry) and tundra (cold‐humid). Here, we test these hypotheses in swallowtail butterflies, a clade with more than 570 species, covering all the continents but Antarctica, and all climatic conditions. Swallowtail butterflies are among the most studied insects, and they are a model group for evolutionary biology and ecology studies. Continental macroecological rules are normally tested using vertebrates, this means that there are fewer examples exploring terrestrial invertebrate patterns at global scale. Here, we compiled a large Geographic Information System database on swallowtail butterflies' distribution maps and used the most complete time‐calibrated phylogeny to quantify diversification rates (DRs). In this paper, we aim to answer the following questions: (1) Are there more biome‐specialist swallowtail butterflies than biome generalists? (2) Is DR related to biome specialization? (3) If so, do swallowtail butterflies inhabiting extreme biomes show higher DRs? (4) What is the effect of species distribution area? Our results showed that swallowtail family presents a great number of biome specialists which showed substantially higher DRs compared to generalists. We also found that biome specialists are unevenly distributed across biomes. Overall, our results are consistent with the resource‐use hypothesis, species climatic niche and biome fragmentation as key factors promoting isolation.

Fossils Do Not Substantially Improve, and May Even Harm, Estimates of Diversification Rate Heterogeneity.

The fossilized birth-death (FBD) model is a naturally appealing way of directly incorporating fossil information when estimating diversification rates. However, an important yet often overlooked property of the original FBD derivation is that it distinguishes between two types of sampled lineages. Here, we first discuss and demonstrate the impact of severely undersampling, and even not including fossils that represent samples of lineages that also had sampled descendants. We then explore the benefits of including fossils, generally, by implementing and then testing two types of FBD models, including one that converts a fossil set into stratigraphic ranges, in more complex likelihood-based models that assume multiple rate classes across the tree. Under various simulation scenarios, including a scenario that exists far outside the set of models we evaluated, including fossils rarely outperform analyses that exclude them altogether. At best, the inclusion of fossils improves precision but does not influence bias. Similarly, we found that converting the fossil set to stratigraphic ranges, which is one way to remedy the effects of undercounting the number of k-type fossils, results in turnover rates and extinction fraction estimates that are generally underestimated. Although fossils remain essential for understanding diversification through time, in the specific case of understanding diversification given an existing, largely modern tree, they are not especially beneficial. [Fossilized birth-death; fossils; MiSSE; state speciation extinction; stratigraphic ranges; turnover rate.].

Out of the temperate zone: A phylogenomic test of the biogeographical conservatism hypothesis in a contrarian clade of ants

AbstractAimThe standard latitudinal diversity gradient (LDG), in which species richness decreases from equator to pole, is a pervasive pattern observed in most organisms. Some lineages, however, exhibit inverse LDGs. Seemingly problematic, documenting and studying contrarian groups can advance understanding of LDGs generally. Here, we identify one such contrarian clade and use a historical approach to evaluate alternative hypotheses that might explain the group's atypical diversity pattern. We focus on the biogeographical conservatism hypothesis (BCH) and the diversification rate hypothesis (DRH).LocationGlobal.TaxonAnts (Hymenoptera: Formicidae: Stenammini).MethodsWe examined the shape of the LDG in Stenammini by plotting latitudinal midpoints for all extant, described species. We inferred a robust genome‐scale phylogeny using UCE data. We estimated divergence dates using beast2 and tested several biogeographical models in BioGeoBEARS. To examine diversification rates and test for a correlation between rate and latitude, we used the programs BAMM and STRAPP, respectively.ResultsStenammini has a skewed inverse LDG with a richness peak in the northern temperate zone. Phylogenomic analyses revealed five major clades and several instances of non‐monophyly among genera (Goniomma, Aphaenogaster). Stenammini and all its major lineages arose in the northern temperate zone. The tribe originated ~51 Ma during a climatic optimum and then diversified and dispersed southward as global climate cooled. Stenammini invaded the tropics at least seven times, but these events occurred more recently and were not linked with increased diversification. There is evidence for a diversification rate increase in Holarctic Aphaenogaster + Messor, but we found no significant correlation between latitude and diversification rate generally.Main ConclusionsOur results largely support the BCH as an explanation for the inverse latitudinal gradient in Stenammini. The clade originated in the Holarctic and likely became more diverse there due to center‐of‐origin, time‐for‐speciation and niche conservatism effects, rather than latitudinal differences in diversification rate.

Radiation of mushroom-forming fungi correlates with novel modes of protecting sexual fruiting bodies

The protection of vulnerable developing structures evolved repeatedly in terrestrial organisms and includes, among others, viviparity in animals and the seed in land plants. In mushroom-forming fungi (Agaricomycetes), sexual spores are born on fruiting bodies, the growth of which is a complex developmental process that is exposed to environmental factors (e.g., desiccation, fungivorous animals). Mushroom-forming fungi evolved a series of innovations in fruiting body protection, however, how these emerged is obscure, leaving the evolutionary principles of fruiting body development poorly known. Here, we show that developmental innovations that lead to the spore-producing surface (hymenophore) being enclosed in a protected environment display asymmetry in their evolution and are associated with increased diversification rates. ‘Enclosed’ development evolved convergently and became a dominant developmental type in several clades of mushrooms. This probably mirrors spore production benefits for species with protected fruiting body initials, by better coping with environmental factors. Our observations highlight new morphological traits associated with mushroom diversification that parallel the evolution of protection strategies in other organisms, such as viviparity or the seed in animals or plants, respectively, but in the context of spore development, highlighting the general importance of protecting vulnerable progeny across the tree of life.

Linking evolutionary dynamics to species extinction for flowering plants in global biodiversity hotspots

AbstractAimBiodiversity hotspots are regions with the highest species richness, and the most threatened species. Previous studies have shown that the extinction risk may be more related to evolutionary history than to species' traits. However, there is a knowledge gap on the relationship between evolutionary history and species extinction risk in biodiversity hotspots. Here, we link evolutionary history to species extinction risk for flowering plants (angiosperms) in global biodiversity hotspots.LocationGlobal biodiversity hotspots.MethodsWe calculated historical evolutionary measures (family species richness, family age, family diversification rate and the ratio of crown age to stem age) of angiosperms from 36 global biodiversity hotspots, which were grouped into 27 regions. Bayesian binomial‐logit univariate and multivariable regression models in conjunction with phylogenetic control and null models were used to identify the evolutionary history predictors of extinction risk for the 27 regions as a whole and for each of the regions.ResultsFamily species richness, family age and family diversification rate are all good indicators for predicting extinction risk. Specifically, when all the 27 regions were considered as a whole, families with higher species richness, older age and/or faster diversification rates have a higher risk of species extinction. However, high extinction risk in some regions, especially in temperate regions, tends to occur in families with low species richness and low diversification rates.Main conclusionsGeographic origin and evolutionary history of species should be jointly taken into account in conservation planning. In general, protection priority should be given to families with high species richness, ancient origins and fast diversification rate because these families seem to be prone to higher extinction risk. Additionally, different strategies should be applied to protect biodiversity in temperate versus tropical regions given that factors affecting extinction risk are different between these regions.

A subfunctionalisation model of gene family evolution predicts balanced tree shapes

We consider a subfunctionalisation model of gene family evolution. A family of n genes that perform z functions is represented by an n×z binary matrix Yt where a 1 in the ijth position indicates that gene i can perform function j. Yt evolves according to a continuous time Markov chain (CTMC) that represents the processes of gene duplication, coding region loss and regulatory region loss with the restriction that each function is protected by selection, meaning that each column in the matrix must contain at least one 1.We generate gene trees based on the CTMC {Yt,t⩾0}. We analyse the long-run behaviour of the model and specify the conditions where we expect gene trees to continue to grow and where we expect them to have a stable number of genes. We show that different choices of rate parameters for the processes of duplication and loss lead to different tree shapes as measured by two common tree-shape statistics: the β-statistic for measuring tree balance and the γ-statistic for assessing diversification rate. We use an extension of β that is estimated from sets of trees. This extension is less biased compared to using the average β value for individual trees.When the rate of gene duplication dominates the rate of gene loss, the process is unstable and the distribution of tree shapes is close to following the uniform ranked tree shape (URT) distribution. However, when the process is stable, gene trees are predicted to have positive values of β indicating balanced trees and negative values of γ indicating that diversification occurs more towards the root of the tree. The results of our analyses suggest that comparing the tree-shape statistics of empirical gene-trees to the predictions presented here will provide a test of the subfunctionalisation model.

Relict lineages with extreme ecology and physiology: metal hyperaccumulation on ultramafic substrates in New Caledonian Alseuosmineae (Asterales)

ABSTRACT Background Relict lineages are an important component of biodiversity, but it is unclear under what circumstances these groups persist. A potential example of such a group is the Alseuosmineae (Asterales) of Oceania. This clade contains the three small families – Alseuosmiaceae, Argophyllaceae and Phellinaceae. The clade has highest diversity in New Caledonia, where there are extensive ultramafic substrates, creating an extreme edaphic environment. Aims Using several lines of evidence we aimed to show that Alseuosmineae qualify as a relict lineage, with specialised adaptations for ultramafic substrates. Methods Molecular phylogenetic and dating analyses were carried out on representatives from all Alseuosmineae genera. Metal concentration in 33 out of 44 Alseuosmineae species was measured in herbarium specimens with X-ray fluorescence spectrometry. Results Dating analyses indicated that Alseuosmineae began diversifying about 75–80 million years ago, and had much slower net diversification rates than other Asterales groups. One-third of the species occur on ultramafic substrates in New Caledonia. Six are categorised as nickel hyperaccumulators, and two as manganese hyperaccumulators. Conclusions Extinction was probably paramount in the history of Alseuosmineae, especially for continental species. We postulate that their adaptation to ultramafic substrates and metal accumulation may have contributed to their survival until today.

Exceptions to the rule: Relative roles of time, diversification rates and regional energy in shaping the inverse latitudinal diversity gradient

AbstractAimInverse latitudinal diversity gradients (i‐LDGs), whereby regional richness peaks outside the tropics, have rarely been investigated, and their causes remain unclear. Here, we investigate three prominent explanations, postulating that species‐rich regions have had: (1) longer time to accumulate species; (2) faster diversification; and (3) more energy to support species‐rich communities. These mechanisms have been shown to explain the tropical megadiversity, and we examine whether they can also explain i‐LDG.LocationGlobal.Time periodContemporary.Major taxa studiedAmphibians, birds and mammals.MethodsWe estimated the time for species accumulation, regional diversification rates and regional energy for six tetrapod taxa (c. 800 species). We quantified the relative effects and interactions among these three classes of variables, using variance partitioning, and confirmed the results across alternative metrics for time (community phylometrics and BioGeoBEARS), diversification rates (BAMM and DR) and regional energy (past and current temperature, and productivity).ResultsAlthough regional richness across each of the six taxa peaked in the temperate region, it varied markedly across hemispheres and continents. The effects of time, diversification rates and regional energy varied greatly from one taxon to another, but high diversification rates generally emerged as the best predictor of high regional richness. The effects of time and regional energy were limited, with the exception of salamanders and cetaceans.Main conclusionsTogether, our results indicate that the causes of i‐LDG are highly taxon specific. Consequently, large‐scale richness gradients might not have a universal explanation, and different causal pathways might converge on similar gradients. Moreover, regional diversification rates might vary dramatically between similar environments and, depending on the taxon, regional richness might or might not depend on the time for species accumulation. Collectively, these results underscore the complexity behind the formation of richness gradients, which might involve a symphony of variations on the interplay of time, diversification rates and regional energy.

Phylogenomics Resolves the Phylogeny of Theaceae by Using Low-Copy and Multi-Copy Nuclear Gene Makers and Uncovers a Fast Radiation Event Contributing to Tea Plants Diversity

Simple SummaryThe Theaceae includes more than 300 species of great morphological diversity and has immense economic, cultural, and ornamental values. However, the evolutionary history of this family remains elusive. We integrated 91 genomes and transcriptome datasets of Theaceae and successfully resolved the phylogeny of Theaceae including relatives of cultivated tea plants from both extensive low-copy and multi-copy nuclear gene markers. Bayes-based molecular dating revealed that the ancestor of the tea family originated slightly earlier than the K-Pg boundary (Mass extinction events including the extinction of dinosaurs) with early diversification of three tribes associated with the Early Eocene Climatic Optimum. Further speciation analysis suggested a sole significant diversification shift rate in the common ancestor of Camellia associated with the Mid-Miocene Climatic Optimum. Collectively, polyploidy events, and key morphological innovation characters, such as pericarp with seed coat hardening, could possibly contribute to the Theaceae species diversity.Tea is one of the three most popular nonalcoholic beverages globally and has extremely high economic and cultural value. Currently, the classification, taxonomy, and evolutionary history of the tea family are largely elusive, including phylogeny, divergence, speciation, and diversity. For understanding the evolutionary history and dynamics of species diversity in Theaceae, a robust phylogenetic framework based on 1785 low-copy and 79,103 multi-copy nuclear genes from 91 tea plant genomes and transcriptome datasets had been reconstructed. Our results maximumly supported that the tribes Stewartieae and Gordonieae are successive sister groups to the tribe Theeae from both coalescent and super matrix ML tree analyses. Moreover, in the most evolved tribe, Theeae, the monophyletic genera Pyrenaria, Apterosperma, and Polyspora are the successive sister groups of Camellia. We also yield a well-resolved relationship of Camellia, which contains the vast majority of Theaceae species richness. Molecular dating suggests that Theaceae originated in the late L-Cretaceous, with subsequent early radiation under the Early Eocene Climatic Optimal (EECO) for the three tribes. A diversification rate shift was detected in the common ancestors of Camellia with subsequent acceleration in speciation rate under the climate optimum in the early Miocene. These results provide a phylogenetic framework and new insights into factors that likely have contributed to the survival of Theaceae, especially a successful radiation event of genus Camellia members to subtropic/tropic regions. These novel findings will facilitate the efficient conservation and utilization of germplasm resources for breeding cultivated tea and oil-tea. Collectively, these results provide a foundation for further morphological and functional evolutionary analyses across Theaceae.

Diversity dynamics of microfossils from the Cretaceous to the Neogene show mixed responses to events.

Microfossils have a ubiquitous and well‐studied fossil record with temporally and spatially fluctuating diversity, but how this arises and how major events affect speciation and extinction is uncertain. We present one of the first applications of PyRate to a micropalaeontological global occurrence dataset, reconstructing diversification rates within a Bayesian framework from the Mesozoic to the Neogene in four microfossil groups: planktic foraminiferans, calcareous nannofossils, radiolarians and diatoms. Calcareous and siliceous groups demonstrate opposed but inconsistent responses in diversification. Radiolarian origination increases from c. 104 Ma, maintaining high rates into the Cenozoic. Calcareous microfossil diversification rates significantly declines across the Cretaceous–Palaeogene boundary, while rates in siliceous microfossil groups remain stable until the Paleocene–Eocene transition. Diversification rates in the Cenozoic are largely stable in calcareous groups, whereas the Palaeogene is a turbulent time for diatoms. Diversification fluctuations are driven by climate change and fluctuations in sea surface temperatures, leading to different responses in the groups generating calcareous or siliceous microfossils. Extinctions are apparently induced by changes in anoxia, acidification and stratification; speciation tends to be associated with upwelling, productivity and ocean circulation. These results invite further micropalaeontological quantitative analysis and study of the effects of major transitions in the fossil record. Despite extensive occurrence data, regional diversification events were not recovered; neither were some global events. These unexpected results show the need to consider multiple spatiotemporal levels of diversity and diversification analyses and imply that occurrence datasets of different clades may be more appropriate for testing some hypotheses than others.

A study of the chromosome number and genome size of the rare species Rhododendron keiskei var. hypoglaucum in Korea

Rhododendron keiskei var. hypoglaucum (Ericaceae) was recently reported in Korea, with a disjunct distribution on the southern islands of the Korean Peninsula. Although chromosome numbers and ploidy variations are important traits in angiosperms, gaining a clear understanding the cytological features of Rhododendron has been hampered by the small size of its chromosomes. We herein report the chromosome number, karyotype structure, and genome size of R. keiskei var. hypoglaucum for the first time. The chromosome number of the investigated plants was 2n = 26 with x = 13 as the base chromosome number, which is the one of the frequently encountered base chromosome numbers in Rhododendron. The karyotype of R. keiskei var. hypoglaucum is composed of metacentric and submetacentric chromosomes similar in length, which ranged from 1.39 to 2.40 μm. The DNA 1C-value in all examined accessions was small, ranging from 0.63 to 0.65 pg, further supporting the stable genome size in Rhododendron. These comprehensive cytological results provide a framework for detailed molecular, cytogenetic, and phylogenomic analyses that can be used to interpret the slow species diversification rate in Rhododendron.