Bayesian Relaxed Molecular Clock Research Articles

Divergence time estimates for the hypoxia-inducible factor-1 alpha (HIF1α) reveal an ancient emergence of animals in low-oxygen environments.

Unveiling the tempo and mode of animal evolution is necessary to understand the links between environmental changes and biological innovation. Although the earliest unambiguous metazoan fossils date to the late Ediacaran period, molecular clock estimates agree that the last common ancestor (LCA) of all extant animals emerged ~850 Ma, in the Tonian period, before the oldest evidence for widespread ocean oxygenation at ~635-560 Ma in the Ediacaran period. Metazoans are aerobic organisms, that is, they are dependent on oxygen to survive. In low-oxygen conditions, most animals have an evolutionarily conserved pathway for maintaining oxygen homeostasis that triggers physiological changes in gene expression via the hypoxia-inducible factor (HIFa). However, here we confirm the absence of the characteristic HIFa protein domain responsible for the oxygen sensing of HIFa in sponges and ctenophores, indicating the LCA of metazoans lacked the functional protein domain as well, and so could have maintained their transcription levels unaltered under the very low-oxygen concentrations of their environments. Using Bayesian relaxed molecular clock dating, we inferred that the ancestral gene lineage responsible for HIFa arose in the Mesoproterozoic Era, ~1273 Ma (Credibility Interval 957-1621 Ma), consistent with the idea that important genetic machinery associated with animals evolved much earlier than the LCA of animals. Our data suggest at least two duplication events in the evolutionary history of HIFa, which generated three vertebrate paralogs, products of the two successive whole-genome duplications that occurred in the vertebrate LCA. Overall, our results support the hypothesis of a pre-Tonian emergence of metazoans under low-oxygen conditions, and an increase in oxygen response elements during animal evolution.

2D approach to reconstruct the evolutionary history of clevelandellids (Ciliophora, Armophorea) inhabiting the hindgut of the Panesthiinae cockroaches

AbstractThe ciliate order Clevelandellida unites endosymbionts of the digestive tract of a variety of invertebrates and vertebrates. In the present study, the primary and secondary structures of nuclear and hydrogenosomal rRNA molecules were employed to reconstruct the phylogenetic relationships and to estimate the divergence times of clevelandellids inhabiting the hindgut of the Panesthiinae cockroaches. The secondary structure information was incorporated in phylogenetic analyses using two different strategies, viz., indirectly through 2D‐guided alignments and directly through so‐called pseudo‐protein data. Nuclear and hydrogenosomal markers carried a consistent phylogenetic signal and robustly supported the monophyletic origin of the family Clevelandellidae as well as of its four genera. According to Bayesian relaxed molecular clock analyses, the last common ancestor (LCA) of the family Clevelandellidae very likely emerged during the Late Cretaceous in the Oriental region. Its descendants most likely expanded to Australia in concert with the Neogene colonization and radiation of their host Panesthiinae cockroaches. Taking into account the time‐calibrated phylogenies and the fact that early branching members of the order Clevelandellida inhabit the digestive tracts of amphibians, it is tempting to speculate that the LCA of the Clevelandellida evolved in ectothermic vertebrates. Amphibians could have brought clevelandellids to the land, where they may have been transmitted to the cockroach digestive tract upon feeding on amphibian faeces.

Molecular data suggests a Pliocene – Early Pleistocene divergence date for the common and pygmy hippopotamus

Hippopotamid phylogeny has proven difficult to resolve with proposed relationships between extant and fossil species receiving mixed levels of support. Of particular interest is the divergence between the two extant hippopotamid species, the well-known common hippopotamus (Hippopotamus amphibius) and the enigmatic pygmy hippopotamus (Choeropsis liberiensis). Previous studies have relied on morphological and fossil evidence to identify the ancestral species at the core of this divergence and its timing. In this study, we assembled a molecular matrix of 26 nuclear gene sequences from 11 ungulate species with two primates as an outgroup. We used a Bayesian relaxed molecular clock approach to reconstruct a calibrated time tree for Cetartiodactyla and estimate the divergence date between the common and pygmy hippopotamus. While previous morphological studies have estimated this event to have occurred sometime during the Late Miocene (between 11.6 and 5.3 million years ago), our nuclear gene-based estimates suggest a more recent split of about 4.04 Ma (95% confidence interval: 8.31–1.97 Ma) via RelTime-ML or 2.4 Ma (95% confidence interval: 3.1–1.6 Ma) via MCMCTree. These more recent estimates correspond with the Early Pliocene – Early Pleistocene sub-epochs and align most closely with the results of previous genomic studies. We discuss how our results compare with previous estimates based on both morphological and molecular studies, some of which extend the predicted range of this divergence date even further back in time. Our results suggest a different path of evolution for the understudied pygmy hippopotamus and reveal that morphological evidence alone may not resolve the correct hippopotamid phylogenetic and time trees. We suggest that the common and pygmy hippopotamus may be phylogenetically closer than once believed. Our results also call for further studies to develop a combined approach incorporating both molecular and morphological evidence to reach a consensus on the evolutionary patterns and timing that led to modern hippopotamid evolution.

Phylogeography of the mesic-adapted striped mouse, Rhabdomys dilectus chakae (Rodentia: Muridae) in forest margins of the Eastern Cape and southern KwaZulu-Natal provinces of South Africa

Abstract Rhabdomys is a genus that occupies a variety of habitats, including forest margins. Among the Rhabdomys taxa, Rhabdomys dilectus chakae has a distribution that covers the eastern seaboard of South Africa, with a poorly defined divergence date from its sister taxon Rhabdomys dilectus dilectus. Here, we study three mitochondrial markers (cytochrome b, cytochrome c oxidase I and partial control region) of R. d. chakae across the Eastern Cape and KwaZulu-Natal forests of South Africa, aiming to determine the cladogenesis (molecular dating) and effective population size of this subspecies through time, in addition to its cladogenesis in relationship to other species within the genus. A phylogenetic analysis revealed six clades within Rhabdomys, confirming that our study area is occupied solely by R. d. chakae, to the exclusion of other recognized sibling species. A fossil-calibrated Bayesian relaxed molecular clock estimated a recent split between R. d. chakae and R. d. dilectus ~1.4 ± 0.35 Mya and between two Rhabdomys pumilio groups, coastal A and B, at 1.16 ± 0.44 Mya. Coalescent Bayesian skyline plots revealed a stable population of R. d. chakae in the study area that was in slow decline until 2500 years ago, when there was an expansion in the late Holocene. Radiation within Rhabdomys dates as far back as 4.27 Mya, and subsequent demographic fluctuations primarily reflect palaeoclimatic changes.

Assembly of a hybrid mangrove, Bruguiera hainesii, and its two ancestral contributors, Bruguiera cylindrica and Bruguiera gymnorhiza

Mangroves are plants that live in tropical and subtropical coastal regions of the world, they are adapted to high salt environments and cyclic tidal flooding. Mangroves play important ecological roles, including acting as breeding grounds for many fish species and to prevent coastal erosion. The genomes of three mangrove species, Bruguiera gymnorhiza, Bruguiera cylindrica, and a hybrid of the two, Bruguiera hainesii were sequenced, assembled and annotated. The two progenitor species, B. gymnorhiza and B. cylindrica, were found to be highly similar to each other and sufficiently similar to B. parviflora to allow it to be used for reference based scaffolding to generate chromosome level scaffolds. The two subgenomes of B. hainesii were independently assembled and scaffolded. Analysis of B. hainesii confirms that it is a hybrid and the hybridisation event was estimated at 2.4 to 3.5 million years ago using a Bayesian Relaxed Molecular Clock approach.

Western Tethys origin, tropical Asia and tropical America disjunction in Berchemia and reinstatement of Phyllogeiton (Rhamneae, Rhamnaceae)

AbstractDespite the well‐known biogeographic pattern in the Northern Hemisphere of disjunct distributions in plants between eastern Asia and North America, many details regarding the evolution of the disjunction between lineages of tropical Asia and tropical America remain poorly understood. Berchemia (Rhamnaceae), comprising ca. 32 species, is distributed predominantly in tropical regions of east to southeast Asia and North America. However, the phylogeny and biogeography of this genus are still not clear. Furthermore, fossils of the genus have been found in Europe, making it an ideal group to understand the biogeographical story of tropical disjunction in the Northern Hemisphere. With two nuclear (ETS, ITS) and three plastid (petA‐psbJ, psbA‐trnH, trnL‐F) DNA regions, we first reconstructed the phylogeny of Berchemia with Bayesian inference and maximum likelihood approaches. Our results indicated that Berchemia, excluding the African species B. discolor and B. zeyheri, which were placed into the reinstated Phyllogeiton, was well‐supported as monophyletic. Divergence times and ancestral areas were inferred using a Bayesian uncorrelated lognormal relaxed molecular clock model and Bayesian binary MCMC method, respectively. The core Berchemia likely originated in the Western Tethys during the middle Eocene, then formed the tropical Asia and tropical America disjunction during the Oligocene and further diversified in Asia from the early Miocene on. The formation and diversification of the core Berchemia likely resulted from a mixture of geologic events, such as the close of the Tethys Ocean, disappearance of the North Atlantic Land Bridges, uplift of the Qinghai‐Tibetan Plateau, and climatic events, such as the establishment and enhancement of Asian monsoons.

Tropical Asian Origin, boreotropical migration and long-distance dispersal in Nettles (Urticeae, Urticaceae)

The tribe Urticeae (Urticaceae), popularly known as Nettles, include 12 genera and ca. 200 species, constituting a diverse and cosmopolitan plant clade centered in tropical Asia, Africa, and South America. The global distribution of this clade makes it an excellent group to test hypotheses regarding the processes underlying tropical intercontinental disjunctions. More specifically, it allows us to test whether current distribution patterns resulted from recent transoceanic long-distance dispersal or ancient vicariance after boreotropical migration. We reconstructed the phylogeny of Nettles with the nuclear ITS and four plastid DNA regions (rbcL, trnL-F, matK and rpl14-rpl36) using Bayesian inference and maximum parsimony approaches. We inferred divergence times using a Bayesian uncorrelated lognormal relaxed molecular clock model and ancestral areas using the divergence-extinction-cladogenesis (DEC) model. Our results indicate a tropical Asian origin for the tribe during the late Paleocene. Migration events to Eurasia, South America and Africa occurred mainly during the Oligocene and Miocene. However, several long-distance dispersal events, including dispersals from Asia to Hawaii or Australasia, were inferred to have occurred from the Miocene onwards. The fleshy fruits and winged diaspores of several taxa are suited for long-distance dispersal.

Species Diversity, Phylogeny, Divergence Time, and Biogeography of the Genus Sanghuangporus (Basidiomycota).

“Sanghuang” is a popular fungus used as a Chinese traditional medicine. In fact, it represents a group of fungi belonging to the genus Sanghuangporus, but little is known about its origin and biogeography. The aim of this study was to characterize the molecular relationships, origin and biogeographical distribution of Sanghuangporus. The multi-locus phylogenetic analyses were used to infer the phylogenetic relationships. In addition, based on Bayesian evolutionary analysis using sequences from the internal transcribed spacer (ITS), nuclear large subunit rDNA (nLSU), translation elongation factor 1-α (EF1-α), and the largest and second largest subunits of RNA polymerase II (RPB1 and RPB2), we used a fungus fossil-based approach to gain insight into the divergence time of species in Sanghuangporus. The molecular phylogeny strongly supports the monophyly of Sanghuangporus (MP = 100%, ML = 100%, and BPP = 1.00), and 13 species are recognized in this genus. The Bayesian uncorrelated lognormal relaxed molecular clock using BEAST and reconstructed ancestral areas indicate that the maximum crown age of Sanghuangporus is approximately 30.85 million years. East Asia is the likely ancestral area (38%). Dispersal and differentiation to other continents then occurred during the late Middle Miocene and Pliocene. The ancestor of Sanghuangporus probably originated in palaeotropical Northeast Asia and covered Northeast Asia and East Africa during the Oligocene-Miocene, hosted by plants that expanded via the “Gomphotherium Landbridge.” Six kinds of dispersal routes are proposed, including intercontinental dispersal events of three clades between Northeast Asia and East Africa, between East Asia and North America, and between Northeast Asia and Europe.

Cenozoic aridization in Central Eurasia shaped diversification of toad-headed agamas (Phrynocephalus; Agamidae, Reptilia).

We hypothesize the phylogenetic relationships of the agamid genus Phrynocephalus to assess how past environmental changes shaped the evolutionary and biogeographic history of these lizards and especially the impact of paleogeography and climatic factors. Phrynocephalus is one of the most diverse and taxonomically confusing lizard genera. As a key element of Palearctic deserts, it serves as a promising model for studies of historical biogeography and formation of arid habitats in Eurasia. We used 51 samples representing 33 of 40 recognized species of Phrynocephalus covering all major areas of the genus. Molecular data included four mtDNA (COI, ND2, ND4, Cytb; 2,703 bp) and four nuDNA protein-coding genes (RAG1, BDNF, AKAP9, NKTR; 4,188 bp). AU-tests were implemented to test for significant differences between mtDNA- and nuDNA-based topologies. A time-calibrated phylogeny was estimated using a Bayesian relaxed molecular clock with nine fossil calibrations. We reconstructed the ancestral area of origin, biogeographic scenarios, body size, and the evolution of habitat preference. Phylogenetic analyses of nuDNA genes recovered a well-resolved and supported topology. Analyses detected significant discordance with the less-supported mtDNA genealogy. The position of Phrynocephalus mystaceus conflicted greatly between the two datasets. MtDNA introgression due to ancient hybridization best explained this result. Monophyletic Phrynocephalus contained three main clades: (I) oviparous species from south-western and Middle Asia; (II) viviparous species of Qinghai–Tibetan Plateau (QTP); and (III) oviparous species of the Caspian Basin, Middle and Central Asia. Phrynocephalus originated in late Oligocene (26.9 Ma) and modern species diversified during the middle Miocene (14.8–13.5 Ma). The reconstruction of ancestral areas indicated that Phrynocephalus originated in Middle East–southern Middle Asia. Body size miniaturization likely occurred early in the history of Phrynocephalus. The common ancestor of Phrynocephalus probably preferred sandy substrates with the inclusion of clay or gravel. The time of Agaminae radiation and origin of Phrynocephalus in the late Oligocene significantly precedes the landbridge between Afro-Arabia and Eurasia in the Early Miocene. Diversification of Phrynocephalus coincides well with the mid-Miocene climatic transition when a rapid cooling of climate drove progressing aridification and the Paratethys salinity crisis. These factors likely triggered the spreading of desert habitats in Central Eurasia, which Phrynocephalus occupied. The origin of the viviparous Tibetan clade has been associated traditionally with uplifting of the QTP; however, further studies are needed to confirm this. Progressing late Miocene aridification, the decrease of the Paratethys Basin, orogenesis, and Plio–Pleistocene climate oscillations likely promoted further diversification within Phrynocephalus. We discuss Phrynocephalus taxonomy in scope of the new analyses.

The timescale of early land plant evolution

Establishing the timescale of early land plant evolution is essential for testing hypotheses on the coevolution of land plants and Earth's System. The sparseness of early land plant megafossils and stratigraphic controls on their distribution make the fossil record an unreliable guide, leaving only the molecular clock. However, the application of molecular clock methodology is challenged by the current impasse in attempts to resolve the evolutionary relationships among the living bryophytes and tracheophytes. Here, we establish a timescale for early land plant evolution that integrates over topological uncertainty by exploring the impact of competing hypotheses on bryophyte-tracheophyte relationships, among other variables, on divergence time estimation. We codify 37 fossil calibrations for Viridiplantae following best practice. We apply these calibrations in a Bayesian relaxed molecular clock analysis of a phylogenomic dataset encompassing the diversity of Embryophyta and their relatives within Viridiplantae. Topology and dataset sizes have little impact on age estimates, with greater differences among alternative clock models and calibration strategies. For all analyses, a Cambrian origin of Embryophyta is recovered with highest probability. The estimated ages for crown tracheophytes range from Late Ordovician to late Silurian. This timescale implies an early establishment of terrestrial ecosystems by land plants that is in close accord with recent estimates for the origin of terrestrial animal lineages. Biogeochemical models that are constrained by the fossil record of early land plants, or attempt to explain their impact, must consider the implications of a much earlier, middle Cambrian-Early Ordovician, origin.

A biogeographic and ecological perspective to the evolution of reproductive behaviour in the family Salamandridae

Amphibians have a complex reproductive behaviour, which shows the highest diversity among tetrapodes. The family Salamandridae, distributed across the entire Holarctic, is one of the most diverse groups of extant salamanders comprising 114 species in 21 genera. The family has a remarkable diversity of courtship modes, amplexus and sperm transfer. It is often hypothesised that this diversity has evolved in adaptation to a specific mating and/or breeding habitat. We test this hypothesis based upon a phylogenetic reconstruction using the complete mitochondrial genome sequences of 45 Salamandridae species, representing all existing genera. We used ancestral character state reconstruction methods and geographic range models and applied relaxed Bayesian molecular clock models to discuss the results in a temporal framework of Salamandridae evolution. Our results show that the family Salamandridae started to diversify in the Late Cretaceous (ca. 87 mya) and is of Western Palearctic origin. Ancestral character state reconstruction predicts that its common ancestor was oviparous, mated on land without amplexus and probably showed a pin wheel spermatophore transfer, which is still found in the Italian endemic Salamandrina terdigidata. Our results suggest that several colonization of continents with subsequent radiations took place, once to the Nearctic and twice into Eastern Asian realms. However, these events were only in one case associated with a change in mating behaviour (dorsal amplexus in Nearctic newts). Around the Cretaceous-Paleogene boundary (K-Pg boundary) several Salamandridae lineages further diverged, again with no obvious changes in mating behaviour. Overall, there is no significant signal for mating character evolution being caused by changes in habitat type, with only a slight tendency that changes in mating habitat might have led to changes in the type of sperm transfer which in turn was associated with changes in the presence or absence of amplexus.

Role of the Qinghai-Tibetan Plateau uplift in the Northern Hemisphere disjunction: evidence from two herbaceous genera of Rubiaceae

To assess the role of the Qinghai-Tibetan Plateau uplift in shaping the intercontinental disjunction in Northern Hemisphere, we analyzed the origin and diversification within a geological timeframe for two relict herbaceous genera, Theligonum and Kelloggia (Rubiaceae). Phylogenetic relationships within and between Theligonum and Kelloggia as well as their relatives were inferred using five chloroplast markers with parsimony, Bayesian and maximum-likelihood approaches. Migration routes and evolution of these taxa were reconstructed using Bayesian relaxed molecular clock and ancestral area reconstruction. Our results suggest the monophyly of each Theligonum and Kelloggia. Eastern Asian and North American species of Kelloggia diverged at ca.18.52 Mya and the Mediterranean species of Theligonum diverged from eastern Asian taxa at ca.13.73 Mya. Both Kelloggia and Theligonum are Tethyan flora relicts, and their ancestors might have been occurred in warm tropical to subtropical environments along the Tethys coast. The Qinghai-Tibetan Plateau separated the eastern and western Tethyan area may contribute significantly to the disjunct distributions of Theligonum, and the North Atlantic migration appears to be the most likely pathway of expansion of Kelloggia to North America. Our results highlight the importance role of the QTP uplift together with corresponding geological and climatic events in shaping biodiversity and biogeographic distribution in the Northern Hemisphere.

African origin and global distribution patterns: Evidence inferred from phylogenetic and biogeographical analyses of ectomycorrhizal fungal genus Strobilomyces

AbstractAimThe ectomycorrhizal genus Strobilomyces is widely distributed throughout many parts of the world, but its origin, divergence and distribution patterns remain largely unresolved. In this study, we aim to explore the species diversity, distribution and evolutionary patterns of Strobilomyces on a global scale by establishing a general phylogenetic framework with extensive sampling.LocationAfrica, Australasia, East Asia, Europe, North America, Central America and Southeast Asia.MethodsThe genealogical concordance phylogenetic species recognition method was used to delimit phylogenetic species. Divergence times were estimated using a Bayesian uncorrelated lognormal relaxed molecular clock. The ancestral area and host of Strobilomyces were inferred via the programs rasp and mesquite. The change of diversification rate over time was estimated using Ape, Laser and Bammtools software packages.ResultsWe recognize a novel African clade and 49 phylogenetic species with morphological evidence, including 18 new phylogenetic species and 23 previously described ones. Strobilomyces probably originated in Africa, in association with Detarioideae/Phyllanthaceae/Monotoideae during the early Eocene. The dispersal to Southeast Asia can be explained by Wolfe's “Boreotropical migration” hypothesis. East Asia, Australasia, Europe and North/Central America are primarily the recipients of immigrant taxa during the Oligocene or later. A rapid radiation implied by one diversification shift was inferred within Strobilomyces during the Miocene.Main conclusionsAn unexpected phylogenetic species diversity within Strobilomyces was uncovered. The highest diversity, resulting probably from a rapid radiation, was found in East Asia. Dispersal played an important role in the current distribution pattern of Strobilomyces. The Palaeotropical disjunction is explained by species dispersal from Africa to Southeast Asia through boreotropical forests during the early Eocene. Species from the Northern Hemisphere and Australasia are largely derived from immigrant ancestors from Southeast Asia.

Evaluating the Impact of Genomic Data and Priors on Bayesian Estimates of the Angiosperm Evolutionary Timescale.

The evolutionary timescale of angiosperms has long been a key question in biology. Molecular estimates of this timescale have shown considerable variation, being influenced by differences in taxon sampling, gene sampling, fossil calibrations, evolutionary models, and choices of priors. Here, we analyze a data set comprising 76 protein-coding genes from the chloroplast genomes of 195 taxa spanning 86 families, including novel genome sequences for 11 taxa, to evaluate the impact of models, priors, and gene sampling on Bayesian estimates of the angiosperm evolutionary timescale. Using a Bayesian relaxed molecular-clock method, with a core set of 35 minimum and two maximum fossil constraints, we estimated that crown angiosperms arose 221 (251-192) Ma during the Triassic. Based on a range of additional sensitivity and subsampling analyses, we found that our date estimates were generally robust to large changes in the parameters of the birth-death tree prior and of the model of rate variation across branches. We found an exception to this when we implemented fossil calibrations in the form of highly informative gamma priors rather than as uniform priors on node ages. Under all other calibration schemes, including trials of seven maximum age constraints, we consistently found that the earliest divergences of angiosperm clades substantially predate the oldest fossils that can be assigned unequivocally to their crown group. Overall, our results and experiments with genome-scale data suggest that reliable estimates of the angiosperm crown age will require increased taxon sampling, significant methodological changes, and new information from the fossil record. [Angiospermae, chloroplast, genome, molecular dating, Triassic.].

Molecular phylogenetic analysis of nuclear genes suggests a Cenozoic over-water dispersal origin for the Cuban solenodon.

The Cuban solenodon (Solenodon cubanus) is one of the most enigmatic mammals and is an extremely rare species with a distribution limited to a small part of the island of Cuba. Despite its rarity, in 2012 seven individuals of S. cubanus were captured and sampled successfully for DNA analysis, providing new insights into the evolutionary origin of this species and into the origins of the Caribbean fauna, which remain controversial. We conducted molecular phylogenetic analyses of five nuclear genes (Apob, Atp7a, Bdnf, Brca1 and Rag1; total, 4,602 bp) from 35 species of the mammalian order Eulipotyphla. Based on Bayesian relaxed molecular clock analyses, the family Solenodontidae diverged from other eulipotyphlan in the Paleocene, after the bolide impact on the Yucatan Peninsula, and S. cubanus diverged from the Hispaniolan solenodon (S. paradoxus) in the Early Pliocene. The strikingly recent divergence time estimates suggest that S. cubanus and its ancestral lineage originated via over-water dispersal rather than vicariance events, as had previously been hypothesised.

Crown group Oxyphotobacteria postdate the rise of oxygen.

The rise of oxygen ca. 2.3billion years ago (Ga) is the most distinct environmental transition in Earth history. This event was enabled by the evolution of oxygenic photosynthesis in the ancestors of Cyanobacteria. However, long-standing questions concern the evolutionary timing of this metabolism, with conflicting answers spanning more than one billion years. Recently, knowledge of the Cyanobacteria phylum has expanded with the discovery of non-photosynthetic members, including a closely related sister group termed Melainabacteria, with the known oxygenic phototrophs restricted to a clade recently designated Oxyphotobacteria. By integrating genomic data from the Melainabacteria, cross-calibrated Bayesian relaxed molecular clock analyses show that crown group Oxyphotobacteria evolved ca. 2.0 billion years ago (Ga), well after the rise of atmospheric dioxygen. We further estimate the divergence between Oxyphotobacteria and Melainabacteria ca. 2.5-2.6Ga, which-if oxygenic photosynthesis is an evolutionary synapomorphy of the Oxyphotobacteria-marks an upper limit for the origin of oxygenic photosynthesis. Together, these results are consistent with the hypothesis that oxygenic photosynthesis evolved relatively close in time to the rise of oxygen.

Phylogeny, biogeography, and diversification of barn owls (Aves: Strigiformes)

The existence of substantial morphological variation has resulted in the description of numerous subspecies of the cosmopolitan barn owl, Tyto alba. However, preliminary studies have revealed a high degree of genetic variation between Old and New World barn owls, suggesting that the T. alba complex may consist of several species. We present a comprehensive study of its taxonomy and propose a spatiotemporal framework to explain the origin and patterns of dispersal and diversification within these cosmopolitan owls. We used a Bayesian relaxed molecular clock approach to assess the timing of diversification. To evaluate the biogeographical pattern, we considered dispersal in addition to temporal connectivity between areas. Finally, we used ecological niche modelling to evaluate their ecological niches. Our phylogenetic analyses suggest that barn owls of the Old and New World show a high degree of genetic divergence, and the barn owls of South and South-east Asia (Tyto alba stertens and Tyto alba javanica) cluster with the Australian barn owl Tyto delicatula. We propose to treat the T. alba complex as three species: T. alba (Africa, Europe), Tyto furcata (New World), and Tyto javanica (Australasia). The dating analyses indicate that the early divergence among the species of the T. alba complex took place in the Middle Miocene and we hypothesize that a common ancestor of the T. alba complex lived in Africa. A potential scenario suggests that T. alba dispersed to Europe and south-western Asia during the interglacial periods of the Miocene/Pliocene, and dispersed into the New World either via an eastern Asian route or a western north Atlantic one.

Breeding system diversification and evolution in American Poa supersect. Homalopoa (Poaceae: Poeae: Poinae).

Background and Aims Poa subgenus Poa supersect. Homalopoa has diversified extensively in the Americas. Over half of the species in the supersection are diclinous; most of these are from the New World, while a few are from South-East Asia. Diclinism in Homalopoa can be divided into three main types: gynomonoecism, gynodioecism and dioecism. Here the sampling of species of New World Homalopoa is expanded to date its origin and diversification in North and South America and examine the evolution and origin of the breeding system diversity.Methods A total of 124 specimens were included in the matrix, of which 89 are species of Poa supersect. Homalopoa sections Acutifoliae, Anthochloa, Brizoides, Dasypoa, Dioicopoa, Dissanthelium, Homalopoa sensu lato (s.l.), Madropoa and Tovarochloa, and the informal Punapoa group. Bayesian and parsimony analyses were conducted on the data sets based on four markers: the nuclear ribosomal internal tanscribed spacer (ITS) and external transcribed spacer (ETS), and plastid trnT-L and trnL-F. Dating analyses were performed on a reduced Poa matrix and enlarged Poaceae outgroup to utilize fossils as calibration points. A relaxed Bayesian molecular clock method was used.Key Results Hermaphroditism appears to be pleisiomorphic in the monophyletic Poa supersect. Homalopoa, which is suggested to have originated in Eurasia 8·4–4·2 million years ago (Mya). The ancestor of Poa supersect. Homalopoa radiated throughout the New World in the Late Miocene–Early Pliocene, with major lineages originating during the Pliocene to Pleistocene (5–2 Mya). Breeding systems are linked to geographic areas, showing an evolutionary pattern associated with different habitats. At least three major pathways from hermaphroditism to diclinism are inferred in New World Homalopoa: two leading to dioecism, one via gynodioecism in South America and another directly from hermaphroditism in North America, a result that needs to be checked with a broader sampling of diclinous species in North America. A third pathway leads from hermaphroditism to gynomonoecism in Andean species of South America, with strictly pistillate species evolving in the highest altitudes.Conclusions Divergence dating provides a temporal context to the evolution of breeding systems in New World Poa supersect. Homalopoa. The results are consistent with the infrageneric classification in part; monophyletic sections are confirmed, it is proposed to reclassify species of sect. Acutifoliae, Dasypoa and Homalopoa s.l. and it is acknowledged that revision of the infrageneric taxonomy of the gynomonoecious species is needed.

Origin of marine planktonic cyanobacteria.

Marine planktonic cyanobacteria contributed to the widespread oxygenation of the oceans towards the end of the Pre-Cambrian and their evolutionary origin represents a key transition in the geochemical evolution of the Earth surface. Little is known, however, about the evolutionary events that led to the appearance of marine planktonic cyanobacteria. I present here phylogenomic (135 proteins and two ribosomal RNAs), Bayesian relaxed molecular clock (18 proteins, SSU and LSU) and Bayesian stochastic character mapping analyses from 131 cyanobacteria genomes with the aim to unravel key evolutionary steps involved in the origin of marine planktonic cyanobacteria. While filamentous cell types evolved early on at around 2,600–2,300 Mya and likely dominated microbial mats in benthic environments for most of the Proterozoic (2,500–542 Mya), marine planktonic cyanobacteria evolved towards the end of the Proterozoic and early Phanerozoic. Crown groups of modern terrestrial and/or benthic coastal cyanobacteria appeared during the late Paleoproterozoic to early Mesoproterozoic. Decrease in cell diameter and loss of filamentous forms contributed to the evolution of unicellular planktonic lineages during the middle of the Mesoproterozoic (1,600–1,000 Mya) in freshwater environments. This study shows that marine planktonic cyanobacteria evolved from benthic marine and some diverged from freshwater ancestors during the Neoproterozoic (1,000–542 Mya).

Phylogenetic Analysis of HIV-1 CRF65_CPX Reveals Yunnan Province Is Still a Source Contributing to the Spread of HIV-1 in China

Phylogenetic Analysis of HIV-1 CRF65_CPX Reveals Yunnan Province Is Still a Source Contributing to the Spread of HIV-1 in China