Reticulate Evolution

High-throughput sequencing is helping biologists to overcome the difficulties of inferring the phylogenies of recently diverged taxa. The present study analyzes the phylogenetic signal of genomic regions with different inheritance patterns using genome skimming and ddRADseq in a species-rich Andean genus (Diplostephium) and its allies. We analyzed the complete nuclear ribosomal cistron, the complete chloroplast genome, a partial mitochondrial genome, and a nuclear-ddRAD matrix separately with phylogenetic methods. We applied several approaches to understand the causes of incongruence among datasets, including simulations and the detection of introgression using the D-statistic (ABBA-BABA test). We found significant incongruence among the nuclear, chloroplast, and mitochondrial phylogenies. The strong signal of hybridization found by simulations and the D-statistic among genera and inside the main clades of Diplostephium indicate reticulate evolution as a main cause of phylogenetic incongruence. Our results add evidence for a major role of reticulate evolution in events of rapid diversification. Hybridization and introgression confound chloroplast and mitochondrial phylogenies in relation to the species tree as a result of the uniparental inheritance of these genomic regions. Practical implications regarding the prevalence of hybridization are discussed in relation to the phylogenetic method.

Heterosis, Haldane and Bateson-Dobzhansky-Muller effects have been widely documented amongst a range of plants and animals. However, typically these effects are shown by taking parents of known genotype into the laboratory and measuring components of the F1 progeny under laboratory conditions. This leaves in doubt the real significance of such effects in the field. Here we use the well-known colour pattern genotypes of the African monarch or queen (Danaus chrysippus), which also control wing length, to test these effects both in the laboratory and in a contact zone in the field. By measuring the wing lengths in animals of known colour pattern genotype we show clear evidence for all three hybrid effects at the A and BC colour patterning loci, and importantly, that these same effects persist in the same presumptive F1s when measured in hybrid populations in the field. This demonstrates the power of a system in which genotypes can be directly inferred in the field and highlights that a...

Background Explaining contrasting patterns of distribution between related species is crucial for understanding the dynamics of biodiversity. Despite instances where hybridization and whole genome duplication (WGD) can yield detrimental outcomes, a role in facilitating the expansion of distribution range has been proposed. The Vitaceae genus Causonis exhibits great variations in species’ distribution ranges, with most species in the derived lineages having a much wider range than those in the early-diverged lineages. Hybridization and WGD events have been suggested to occur in Causonis based on evidence of phylogenetic discordance. The genus, therefore, provides us with an opportunity to for explore different hybridization and polyploidization modes in lineages with contrasting species’ distribution ranges. However, the evolutionary history of Causonis incorporating potential hybridization and WGD events remains to be explored. Results With plastid and nuclear data from dense sampli...

Capsella bursa-pastoris (Brassicaceae) is one of the most successful tetraploid species in the world. It showed high morphological diversity within Egyptian populations. The morphological investigations of herbarium specimens and fresh collected populations grouped them under three distinguished morphotypes (Lobed "L"; Simple "S" and Lobed-Simple "LS") depending mainly on the basal leaves structure. This high degree of phenotypic variation has received our critical attention. Until recently, the previous studies on C. bursa-pastoris attributed its phenotypic variation to environmental factors. But in Egypt, these three morphotypes were traced in mixed populations along with the species geographical range, so the environmental factors have no influence on their distribution or phenotypic variation. Accordingly, our primary concern in this study was to determine the factors controlling this variation. The cytogenetic studies revealed that the three identi...

Background Explaining contrasting patterns of distribution between related species is crucial for understanding the dynamics of biodiversity. Despite instances where hybridization and whole genome duplication (WGD) can yield detrimental outcomes, a role in facilitating the expansion of distribution range has been proposed. The Vitaceae genus Causonis exhibits great variations in species’ distribution ranges, with most species in the derived lineages having a much wider range than those in the early-diverged lineages. Hybridization and WGD events have been suggested to occur in Causonis based on evidence of phylogenetic discordance. The genus, therefore, provides us with an opportunity to for explore different hybridization and polyploidization modes in lineages with contrasting species’ distribution ranges. However, the evolutionary history of Causonis incorporating potential hybridization and WGD events remains to be explored. Results With plastid and nuclear data from dense sampli...

Recent advances in biomolecular sequencing have revealed the important role that interspecific gene flow has played in genome evolution throughout the Tree of Life. Current and future genomic studies will bring large amounts of genomic sequence to bear upon this topic, and scalable computational methodologies are needed to detect and analyze genomic signatures of interspecific introgression in large-scale datasets. To address the methodological gap, we introduce a new computational framework known as PHiMM (or "fast PhyloNet + Hidden Markov Model"). PHiMM combines inference and learning under a combined model of genetic drift, substitutions, recombination, and gene flow with a coalescent-based approximation technique. We compare the performance of PHiMM against the state of the art using synthetic and empirical genomic sequence data. We find that PHiMM offers better computational runtime and main memory usage by multiple orders of magnitude, while returning comparable inference accuracy. An open-source software implementation of the PHiMM framework and open data are publicly available at .

The genus Ficus L. (Moraceae) is known for its diverse morphology and habit as well as economical, ecological, and pharmaceutical importance. In this study, six Philippine Ficus species were utilized to determine the best barcoding loci among cpDNA (rbcL, trnH-psbA) and nrDNA (ITS) markers on the basis of universality, discriminatory power, and resolution of species. Both trnH-psbA and ITS showed 100% PCR success rate and 67% sequencing success, whereas rbcL exhibited 67% PCR success rate but 100% sequencing success. The trnH-psbA marker performed best in terms of discriminatory power, showing the highest variable informative site (71.37%) and the highest mean interspecific distance (48.32% ± 19.89%). ITS ranked next to trnH-psbA since it was able to generate 100% resolution of species and a comparable mean intraspecific distance score with trnH- psbA. Meanwhile, rbcL failed to resolve any species correctly; thus, it has a 0% resolution of species. We initially recommend trnH-psbA a...

Late Neogene and Quaternary climatic oscillations have greatly shaped the genetic structure of the Mediterranean Basin flora, with mountain plant species tracking warm interglacials/cold glacials by means of altitudinal shifts instead of broad latitudinal ones. Such dynamics may have enhanced population divergence but also secondary contacts. In this paper, we use a case example of subsection Willkommia of Centaurea (comprising three narrowly distributed endemic species, Centaurea gadorensis, C. pulvinata, and C. sagredoi) to test for reticulate evolution and recurrent hybridizations between nearby populations. For this, we combine analyses of genetic diversity and structuring, gene flow and spatial correlation, and ecological niche modeling. Our results support the contention that the current genetic structure of the three species is the result of historical gene flow at sites of secondary contact during the glacial periods, followed by isolation after the retraction of populations to the middle-upper areas of the mountains during the interglacial periods. The extent and direction of the gene flow was determined largely by the location of the populations on mountainsides oriented toward the same valley or toward different valleys, suggesting the intermountain valleys as the areas where secondary contacts occurred.

Inferring the ecological and evolutionary processes underlying lineage and phenotypic diversification is of paramount importance to shed light on the origin of contemporary patterns of biological diversity. However, reconstructing phylogenetic relationships in recent evolutionary radiations represents a major challenge due to the frequent co-occurrence of incomplete lineage sorting and introgression. In this study, we combined high throughput sequence data (ddRADseq), geometric morphometric information, and novel phylogenetic inference methods that explicitly account for gene flow to infer the evolutionary relationships and the timing and mode of diversification in a complex of Ibero-Maghrebian montane grasshoppers of the subgenusDreuxius(genusOmocestus). Our analyses supported the phenotypic distinctiveness of most sister taxa, two events of historical introgression involving lineages at different stages of the diversification continuum, and the recent Pleistocene origin (< 1 Ma...

Evolutionary history is typically portrayed as a branching phylogenetic tree, yet not all evolution proceeds in a purely bifurcating manner. Introgressive hybridization is one process that results in reticulate evolution. Most known examples of genome-wide introgression occur among closely related species with relatively recent common ancestry; however, we present evidence for ancient hybridization and genome-wide introgression between major stem lineages of darters, a species-rich clade of North American freshwater fishes. Previous attempts to resolve the relationships of darters have been confounded by the uncertain phylogenetic resolution of the lineage Allohistium. In this study, we investigate the phylogenomics of darters, specifically the relationships of Allohistium, through analyses of approximately 30,000 RADseq loci sampled from 112 species. Our phylogenetic inferences are based on traditional approaches in combination with strategies that accommodate reticulate evolution....

Allopolyploid plants have long been regarded as possessing genetic advantages under certain circumstances due to the combined effects of their hybrid origins and duplicated genomes. However, the evolutionary consequences of allopolyploidy in lineage diversification remain to be fully understood. Here, we investigate the evolutionary consequences of allopolyploidy using 138 transcriptomic sequences of Gesneriaceae, including 124 newly sequenced, focusing particularly on the largest subtribe Didymocarpinae. We estimated the phylogeny of Gesneriaceae using concatenated and coalescent-based methods based on five different nuclear matrices and 27 plastid genes, focusing on relationships among major clades. To better understand the evolutionary affinities in this family, we applied a range of approaches to characterize the extent and cause of phylogenetic incongruence. We found that extensive conflicts between nuclear and chloroplast genomes and among nuclear genes were caused by both inc...

The Pteris fauriei group (Pteridaceae) has a wide distribution in Eastern Asia and includes 18 species with similar but varied morphology. We collected more than 300 specimens of the P. fauriei group and determined ploidy by flow cytometry and inferred phylogenies by molecular analyses of chloroplast and nuclear DNA markers. Our results reveal a complicated reticulate evolution, consisting of seven parental taxa and 58 hybrids. The large number of hybrid taxa have added significant morphological complexity to the group leading to difficult taxonomic issues. The hybrids generally had broader ranges and more populations than their parental taxa. Genetic combination of different pairs of parental species created divergent phenotypes of hybrids, exhibited by both morphological characteristics and ecological fidelities. Niche novelty could facilitate hybrid speciation. Apogamy is common in this group and potentially contributes to the sustainability of the whole group. We propose that fr...

Evolutionary data has been traditionally modeled via phylogenetic trees; however, branching alone cannot model conflicting phylogenetic signals, so networks are used instead. Ancestral recombination graphs (ARGs) are used to model the evolution of incompatible sets of SNP data, allowing each site to mutate only once. The model often aims to minimize the number of recombinations. Similarly, incompatible cluster data can be represented by a reticulation network that minimizes reticulation events. The ARG literature has traditionally been disjoint from the reticulation network literature. By building on results from the reticulation network literature, we resolve an open question of interest to the ARG community. We explicitly prove that the History Bound, a lower bound on the number of recombinations in an ARG for a binary matrix, which was previously only defined procedurally, is equal to the minimum number of reticulation nodes in a network for the corresponding cluster data. To facilitate the proof, we give an algorithm that constructs this network using intermediate values from the procedural History Bound definition. We then develop a top-down algorithm for computing the History Bound, which has the same worst-case runtime as the known dynamic program, and show that it is likely to run faster in typical cases.

Background and Aims Cardamine occulta (Brassicaceae) is an octoploid weedy species (2n = 8x = 64) originated in Eastern Asia. It has been introduced to other continents including Europe and considered to be an invasive species. Despite its wide distribution, the polyploid origin of C. occulta remained unexplored. The feasibility of comparative chromosome painting (CCP) in crucifers allowed us to elucidate the origin and genome evolution in Cardamine species. We aimed to investigate the genome structure of C. occulta in comparison with its tetraploid (2n = 4x = 32, C. kokaiensis and C. scutata) and octoploid (2n = 8x = 64, C. dentipetala) relatives. Methods Genomic in situ hybridization (GISH) and large-scale CCP were applied to uncover the parental genomes and chromosome composition of the investigated Cardamine species. Key Results All investigated species descended from a common ancestral Cardamine genome (n = 8), structurally resembling the Ancestral Crucifer Karyotype (n = 8), b...

The genus Cardamine is distributed worldwide and is rich in polyploids. Here we review the considerable amount of knowledge accumulated on polyploid species and related diploids over the last twenty years, particularly in the Mediterranean area and neighbouring regions of Central Europe. These studies addressed the taxonomic treatment of polyploids, their origin and evolution, phylogenetic relationships to diploid and other polyploid relatives, and overall morphological, genetic, karyological and ecological patterns. The revealed parentage of polyploids such as C. ×asarifolia, C. ×flexuosa and C. ×schulzii shows the power of recently developed molecular and cytogenetic approaches, which enable us to understand how the polyploid taxa originated, became established and further evolved. It is emphasised that diploid-polyploid species complexes, such as the C. amara and C. pratensis groups, represent true natural laboratories for polyploid studies. Particularly the species complex of C. pratensis contains numerous diploid lineages and polyploid populations of all levels up to dodecaploids, including aneuploids and dysploids, providing excellent possibilities for the application of a wide spectrum of methodical approaches to understand the processes underlying polyploid evolution. Hybridization revealed in this genus occurs both at the homoploid and heteroploid levels and mainly in mountain regions, where parapatric and/or ecologically differentiated close relatives come into contact. These studies document that interspecific gene flow can result in various patterns, such as continuous variation in hybrid swarms (C. pratensis × C. raphanifolia), established hybrid species of significant evolutionary potential (C. ×schulzii, C. ×insueta), or largely sterile hybrids of first generations only and persistence through vegetative propagation (C. ×ferrarii, C. ×enriquei). All of these cases also show how recent hybridization and introgression add to the complexity of reticulated patterns of evolution in Cardamine in the Mediterranean and adjacent areas.

The Pteris fauriei group (Pteridaceae) has a wide distribution in Eastern Asia and includes 18 species with similar but varied morphology. We collected more than 300 specimens of the P. fauriei group and determined ploidy by flow cytometry and inferred phylogenies by molecular analyses of chloroplast and nuclear DNA markers. Our results reveal a complicated reticulate evolution, consisting of seven parental taxa and 58 hybrids. The large number of hybrid taxa have added significant morphological complexity to the group leading to difficult taxonomic issues. The hybrids generally had broader ranges and more populations than their parental taxa. Genetic combination of different pairs of parental species created divergent phenotypes of hybrids, exhibited by both morphological characteristics and ecological fidelities. Niche novelty could facilitate hybrid speciation. Apogamy is common in this group and potentially contributes to the sustainability of the whole group. We propose that fr...

SUMMARY: The name Lavatera columbretensis, recently proposed for a hybrid growing in the Columbretes Archipelago (Castellón, E of Spain), is transferred to Malva. Additional data on nomenclature of other related taxa of ‘L. sect. Anthema ’ and ‘L. sect. Axolopha’, are also reported. Key words: Lavatera, Malva, hybrids, Mediter-ranean islands, Spain. RESUMEN: Se transfiere al género Malva el nombre Lavatera columbretensis, recientemente descrito para un híbrido encontrado en el archipiélago de las Columbre-tes (Castellón). Además, se presentan datos complementarios sobre la nomenclatura de otros táxones de ‘L. sect. Anthema ’ y ‘L. sect. Axolopha’, con él relacionados. Pala-bras clave: Lavatera, Malva, híbridos, islas mediterráneas, España.

Puddle frogs of the Phrynobatrachus steindachneri species complex are a useful group for investigating speciation and phylogeography in Afromontane forests of the Cameroon Volcanic Line, western Central Africa. The species complex is represented by six morphologically relatively cryptic mitochondrial DNA lineages, only two of which are distinguished at the species levelsouthern P. jimzimkusi and Lake Oku endemic P. njiomock, leaving the remaining four lineages identified as 'P. steindachneri'. In this study, the six mtDNA lineages are subjected to genomic sequence capture analyses and morphological examination to delimit species and to study biogeography. The nuclear DNA data (387 loci; 571,936 aligned base pairs) distinguished all six mtDNA lineages, but the topological pattern and divergence depths supported only four main clades: P. jimzimkusi, P. njiomock, and only two divergent evolutionary lineages within the four 'P. steindachneri' mtDNA lineages. One of the two lineages is herein described as a new species, P. amieti sp. nov. Reticulate evolution (hybridization) was detected within the species complex with morphologically intermediate hybrid individuals placed between the parental species in phylogenomic analyses, forming a ladder-like phylogenetic pattern. The presence of hybrids is undesirable in standard phylogenetic analyses but is essential and beneficial in the network multispecies coalescent. This latter approach provided insight into the reticulate evolutionary history of these endemic frogs. Introgressions likely occurred during the Middle and Late Pleistocene climatic oscillations, due to the cyclic connections (likely dominating during cold glacials) and separations (during warm interglacials) of montane forests. The genomic phylogeographic pattern supports the separation of the southern (Mt. Manengouba to Mt. Oku) and northern mountains at the onset of the Pleistocene. Further subdivisions occurred in the Early Pleistocene, separating populations from the northernmost (Tchabal Mbabo, Gotel Mts.) and middle mountains (Mt. Mbam, Mt. Oku, Mambilla Plateau), as well as the microendemic lineage restricted to Lake Oku (Mt. Oku). This unique model system is highly threatened as all the species within the complex have exhibited severe population declines in the past decade, placing them on the brink of extinction. In addition, Mount Oku is identified to be of particular conservation importance because it harbors three species of this complex. We, therefore, urge for conservation actions in the Cameroon Highlands to preserve their diversity before it is too late.

Allopolyploidy is considered a driver of diversity in subtribe Loliinae. We investigate the evolution and systematics of the poorly studied Mesoamerican and South American polyploid broad-leaved Festuca L. species of uncertain origin and unclear taxonomy. A taxonomic study of seven diagnostic morphological traits was conducted on a representation of 22 species. Phylogenomic analyses were performed on a representation of these supraspecific taxa and all other Loliinae lineages using separate data from the entire plastome, nuclear rDNA 45S and 5S genes, and repetitive DNA elements. F. subgen. Mallopetalon falls within the fine-leaved (FL) Loliinae clade, whereas the remaining taxa are nested within the broad-leaved (BL) Loliinae clade forming two separate Mexico-Central-South American (MCSAI, MCSAII) lineages. MCSAI includes representatives of F. sect. Glabricarpae and F. subgen. Asperifolia plus F. superba, and MCSAII of F. subgen. Erosiflorae and F. sect. Ruprechtia plus F. argentina. MCSAII likely had a BL Leucopoa paternal ancestor, MCSAI and MCSAII a BL Meso-South American maternal ancestor, and Mallopetalon FL, American I-II ancestors. Plastome vs. nuclear topological discordances corroborated the hybrid allopolyploid origins of these taxa, some of which probably originated from Northern Hemisphere ancestors. The observed data indicate rapid reticulate radiations in the Central-South American subcontinent. Our systematic study supports the reclassification of some studied taxa in different supraspecific Festuca ranks.

While it is known that whole genome duplication (WGD) and reticulate evolution play important roles in plant evolution, the origins and evolutionary histories of most polyploid and reticulate groups are still poorly known. The North Temperate haplostemonous (NTH) Ludwigia L. (sections Isnardia (L.) W. L. Wagner & Hoch, Ludwigia, Microcarpium Munz, and Miquelia P. H. Raven) group, characterized by having 4-merous and haplostemonous flowers, pluriseriate and free seeds, glabrous and convex nectaries, and a north-temperate distribution, is a polyploid complex (23, 43, 63, and 83) of 24 species with frequent reports of inter-and intrasectional hybridization. Although earlier biosystematics studies postulated some evolutionary scenarios and recent molecular phylogenetic studies have partially tested these propositions, the full history of their reticulate evolution remains puzzling. In this study, we sequenced four chloroplast regions (rpL16, rpoB-trnC, trnL-trnF, and ycf6-psbM) and conducted extensive molecular cloning of the biparentally inherited single-copy nuclear PgiC gene (376 clones in total), sampling 23 of the 24 NTH Ludwigia species whose chromosome numbers and ploidy levels were confirmed. Both the chloroplast and PgiC trees include strongly supported sister clades of section Ludwigia (four diploid species) and the "Microcarpium complex" (composed of sections Isnardia, Microcarpium, and Miquelia), which together are sister to the rest of Ludwigia. In the PgiC tree, eight clades are identified within the Microcarpium complex, with four clades including no extant diploid species. Neither sections Isnardia nor Microcarpium are monophyletic, while the monospecific section Miquelia has a hybrid origin. By integrating our phylogenetic trees with previous cytological hypotheses, the reticulate evolution of NTH Ludwigia is disentangled and four to eight extinct diploid species are inferred. Ancestral area reconstruction supports a North American origin of L. ovalis whose current East Asian distribution reflects a relict of the Arcto-Tertiary Geoflora. Based on our results, we propose to synonymize sections Microcarpium and Miquelia under the expanded section Isnardia.

The rapidly increasing number of available plant genomes opens up almost unlimited prospects for biology in general and molecular phylogenetics in particular. A recent study took advantage of this data and identified a set of nuclear genes that occur in single copy in multiple sequenced angiosperms. The present study is the first to apply genomic sequence of one of these low copy genes, agt1, as a phylogenetic marker for specieslevel phylogenetics. Its utility is compared to the performance of several coding and non-coding chloroplast loci that have been suggested as most applicable for this taxonomic level. As a model group, we chose Tildenia, a subgenus of Peperomia (Piperaceae), one of the largest plant genera. Relationships are particularly difficult to resolve within these species rich groups due to low levels of polymorphisms and fast or recent radiation. Therefore, Tildenia is a perfect test case for applying new phylogenetic tools. Results: We show that the nuclear marker agt1, and in particular the agt1 introns, provide a significantly increased phylogenetic signal compared to chloroplast markers commonly used for low level phylogenetics. 25% of aligned characters from agt1 intron sequence are parsimony informative. In comparison, the introns and spacer of several common chloroplast markers (trnK intron, trnK-psbA spacer, ndhF-rpl32 spacer, rpl32-trnL spacer, psbA-trnH spacer) provide less than 10% parsimony informative characters. The agt1 dataset provides a deeper resolution than the chloroplast markers in Tildenia. Conclusions: Single (or very low) copy nuclear genes are of immense value in plant phylogenetics. Compared to other nuclear genes that are members of gene families of all sizes, lab effort, such as cloning, can be kept to a minimum. They also provide regions with different phylogenetic content deriving from coding and non-coding parts of different length. Thus, they can be applied to a wide range of taxonomic levels from family down to population level. As more plant genomes are sequenced, we will obtain increasingly precise information about which genes return to single copy most rapidly following gene duplication and may be most useful across a wide range of plant groups.

Understanding the factors that cause heterogeneity among gene trees can increase the accuracy of species trees. Discordant signals across the genome are commonly produced by incomplete lineage sorting (ILS) and introgression, which in turn can result in reticulate evolution. Species tree inference using the multispecies coalescent is designed to deal with ILS and is robust to low levels of introgression, but extensive introgression violates the fundamental assumption that relationships are strictly bifurcating. In this study, we explore the phylogenomics of the iconic Liolaemus subgenus of South American lizards, a group of over 100 species mostly distributed in and around the Andes mountains. Using mitochondrial DNA (mtDNA) and genome-wide restriction site-associated DNA sequencing (RADseq; nDNA hereafter), we inferred a time-calibrated mtDNA gene tree, nDNA species trees, and phylogenetic networks. We found high levels of discordance between mtDNA and nDNA, which we attribute in p...

The chiropteran fauna of the island of Saint Vincent, represented by 12 species, is among the most complex in the Lesser Antilles, being represented by four families including Noctilionidae (1 species), Mormoopidae (1), Phyllostomidae (8), and Molossidae (2). This fauna includes four trophic guilds as represented by Noctilio leporinus (piscivore/insectivore); Glossophaga longirostris and Monophyllus plethodon (nectarivore/pollenivore); Artibeus lituratus, A. schwartzi, Brachyphylla cavernarum, Ardops nichollsi, and Sturnira paulsoni (frugivore); and Pteronotus fuscus, Micronycteris buriri, Molossus molossus, and Tadarida brasiliensis (insectivore). One species—Micronycteris buriri—and two subspecies—Sturnira paulsoni paulsoni and Ardops nichollsi vincentensis—are endemic to the island. Recent advancements in population genomics have led to the discovery of the reticulated evolutionary history of Artibeus schwartzi and it is likely that the formation of the hybrid evolutionary trajec...

Also known as the Himalayan Poppy, Meconopsis is a genus of herbaceous plants growing only in the high elevation habits of the Himalaya and its adjacent plateau and mountain areas. The genus exhibits high morphological and ecological diversity, but the major causes of divergence in Meconopsis have not previously been studied. Our recent revised taxonomic classification, based on a molecular phylogeny, divided the genus into four monophyletic sections. Because chromosome number varies among these sections and our previous phylogenetic analyses revealed extensive incongruence between the recovered nrITS and cpDNA trees, possibly due to ancient hybridization, this study focused on evaluating the potential role of ancient polyploidization and hybridization in Meconopsis' evolutionary history. Our investigation based on the results of reconstructed ancestral chromosome numbers using a Maximum Likelihood method implemented in chromEvol showed that two extant Meconopsis sections (sect. Grandes and sect. Primulinae) shared a triploid ancestor. We further examined the pattern of hybridization in Meconopsis by reconstructing a nuclear marker GAPDH (glyceraldehyde 3-phosphate dehydrogenase gene) network. The result, along with morphological, phylogenetic, and cytological evidence, all point to a hybrid nature of the triploid ancestor. Based on the resultant GAPDH network, an ancient reticulate evolution scenario in Meconopsis is proposed. Overall, this preliminary study shows how an ancient triploid event promoted polyploid evolution in Meconopsis and also exemplifies how allotriploidization and successive polyploidization played an important role in diversification of the genus.

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The Iberian mountain spiny fescues are a reticulate group of five diploid grass taxa consisting of three parental species and two putative hybrids: F. × souliei (F. eskia × F. quadriflora) and F. × picoeuropeana (F. eskia × F. gautieri). Phenotypic and molecular studies were conducted with the aim of determining the taxonomic boundaries and genetic relationships of the five taxa and disentangling the origins of the two hybrids. Statistical analyses of 31 selected phenotypic traits were conducted on individuals from 159 populations and on nine type specimens. Molecular analyses of random amplified polymorphic DNA (RAPD) markers were performed on 29 populations. The phenotypic analyses detected significant differences between the five taxa and demonstrated the overall intermediacy of the F. × picoeuropeana and F. × souliei between their respective parents. The RAPD analysis corroborated the genetic differentiation of F. eskia, F. gautieri and F. quadriflora and the intermediate nature of the two hybrids; however, they also detected genetic variation within F. × picoeuropeana. These results suggest distinct origins for F. × picoeuropeana in the Cantabrian and Pyrenean mountains, with the sporadic Pyrenean populations having potentially resulted from recent hybridizations and the stabilized Cantabrian ones from older events followed by potential displacements of the parents.

Stellingen 1. Based upon life history strategy (LHS) studies, Gigasporaceae are "K" strategists in contrast to single sporeproducing "Glomus" species (this thesis). These propositions are part of the Ph.D. thesis "Biology, ecology and evolution of the family Gigasporaceae, Arbuscular Mycorrhizal Fungi (Glomeromycota)" by Francisco Adriano de Souza Leiden,

Biotechnology and information technology are the two generic technologies which are poised to have a major impact in the coming decades. Students of generic technologies have pointed out that their impact would be much greater than the earlier generic technologies associated with the Industrial Revolution and its aftermath, including such technologies as those asssociated with steam, electricity, chemicals and oil1. But the two technologies replace and/or intimately affect respectively life processes and cultural ...

This article shows how to fit reticulate finite and infinite sites sequence spectra to aligned data from five modern human genomes (San, Yoruba, French, Han and Papuan) plus two archaic humans (Denisovan and Neanderthal), to better infer demographic parameters. These include interbreeding between distinct lineages. Major improvements in the fit of the sequence spectrum are made with successively more complicated models. Findings include some evidence of a male biased gene flow from the Denisova lineage to Papuan ancestors and possibly even more archaic gene flow. It is unclear if there is evidence for more than one Neanderthal interbreeding, as the evidence suggesting this largely disappears when a finite sites model is fitted.

Recent advances in understanding CD4(+) T-cell differentiation suggest that previous models of a few distinct, stable effector phenotypes were too simplistic. Although several well-characterized phenotypes are still recognized, some states display plasticity, and intermediate phenotypes exist. As a framework for reexamining these concepts, we use Waddington's landscape paradigm, augmented with explicit consideration of stochastic variations. Our animation program "LAVA" visualizes T-cell differentiation as cells moving across a landscape of hills and valleys, leading to attractor basins representing stable or semistable differentiation states. The model illustrates several principles, including: (i) cell populations may behave more predictably than individual cells; (ii) analogous to reticulate evolution, differentiation may proceed through a network of interconnected states, rather than a single well-defined pathway; (iii) relatively minor changes in the barriers betw...

Natural hybridization between closely related species in sympatry is an evolutionary process that is common in orchids. Once seen as a threat to parent species, interspecific genetic change is increasingly viewed as a source of novel variation in some ecological contexts. Terrestrial fringed orchids in the genus Platanthera contain several clades with high genetic compatibility among species and many putative hybrids. We used biallelic SNPs generated with 3RAD sequencing to study the hybrid complex formed from the parent species P. blephariglottis, P. ciliaris, and P. cristata with high resolution. The genetic structure and phylogenetic relationship of the hybrid complex revealed site-dependent gene flow between species. We documented extensive hybridization and cryptic hybrids in sympatric sites. Interspecific genetic exchange is particularly common between P. blephariglottis and P. ciliaris, with cryptic hybrids among putative P. ciliaris samples being more common than parental as...

The identification of homeologous genomes and the biogeographical analyses of highly reticulate allopolyploid-rich groups face the challenge of incorrectly inferring the genomic origins and the biogeographical patterns of the polyploids from unreliable strictly bifurcating trees. Here we reconstruct a plausible evolutionary scenario of the diverging and merging genomes inherited by the diploid and allopolyploid species and cytotypes of the model grass genus Brachypodium. We have identified the ancestral Brachypodium genomes and inferred the paleogeographical ranges for potential hybridization events that originated its allopolyploid taxa. We also constructed a comprehensive phylogeny of Brachypodium from five nuclear and plastid genes using Species Tree Minimum Evolution allele grafting and Species Network analysis. The divergence ages of the lineages were estimated from a consensus maximum clade credibility tree using fossil calibrations, whereas ages of origin of the diploid and allopolyploid species were inferred from coalescence Bayesian methods. The biogeographical events of the genomes were reconstructed using a stratified Dispersal-Extinction-Colonization model with three temporal windows. Our combined Minimum Evolution-coalescence-Bayesian approach allowed us to infer the origins and the identities of the homeologous genomes of the Brachypodium allopolyploids, matching the expected ploidy levels of the hybrids. To date, the current extant progenitor genomes (species) are only known for B. hybridum. Putative ancestral homeologous genome have been inherited by B. mexicanum, ancestral and recent genomes by B. boissieri, and only recently evolved genomes by B. retusum and the core perennial clade allopolyploids (B. phoenicoides, B. pinnatum 4x, B. rupestre 4x). We dissected the complex spatio-temporal evolution of ancestral and recent genomes and have detected successive splitting, dispersal and merging events for dysploid homeologous genomes in diverse geographical scenarios that have led to the current extant taxa. Our data support Mid-Miocene splits of the Holarctic ancestral genomes that preceded the Late Miocene origins of Brachypodium ancestors of the modern diploid species. Successive divergences of the annual B. stacei and B. distachyon diploid genomes were implied to have occurred in the Mediterranean region during the Late Miocene-Pliocene. By contrast, a profusion of splits, range expansions and different genome mergings were inferred for the perennial diploid genomes in the Mediterranean and Eurasian regions, with sporadic colonizations and further mergings in other continents during the Quaternary. A reliable biogeographical scenario was obtained for the Brachypodium genomes and allopolyploids where homeologous genomes split from their respective diploid counterpart lineages in the same ancestral areas, showing similar or distinct dispersals. By contrast, the allopolyploid taxa remained in the same ancestral ranges after hybridization and genome doubling events. Our approach should have utility in deciphering the genomic composition and the historical biogeography of other allopolyploid-rich organismal groups, which are predominant in eukaryotes.

The wild rice gene pool, i.e., AA-genome, in Australia is geographically and genetically distinct from that in Asia. Two distinct taxa are found growing together in northern Australia,Oryza meridionalis(including annual and perennial forms) and anOryza rufipogonlike taxa that have been shown to have a chloroplast genome sequence that is closer to that ofO. meridionalisthan toO. rufipogonfrom Asia. Rare plants of intermediate morphology have been observed in the wild despite a reported reproductive barrier between these two species. We now report the resequencing of plants from 26 populations including both taxa and putative hybrids. A comparison of chloroplast and nuclear genome sequences indicated re-combinations that demonstrated hybridisation in both directions. Individuals with intermediate morphology had high nuclear genome heterozygosity consistent with a hybrid origin. An examination of specific genes (e.g., starch biosynthesis genes) revealed the presence of heterozygotes wi...

Background Interspecific hybrids between S. cerevisiae × S. kudriavzevii have frequently been detected in wine and beer fermentations. Significant physiological differences among parental and hybrid strains under different stress conditions have been evidenced. In this study, we used comparative genome hybridization analysis to evaluate the genome composition of different S. cerevisiae × S. kudriavzevii natural hybrids isolated from wine and beer fermentations to infer their evolutionary origins and to figure out the potential role of common S. kudriavzevii gene fraction present in these hybrids. Results Comparative genomic hybridization (CGH) and ploidy analyses carried out in this study confirmed the presence of individual and differential chromosomal composition patterns for most S. cerevisiae × S. kudriavzevii hybrids from beer and wine. All hybrids share a common set of depleted S. cerevisiae genes, which also are depleted or absent in the wine strains studied so far, and the p...

Several recent studies have suggested that hybridization may play a previously unrecognized and important role in the evolution of corals. Our observations of polymorphic and recombinant sequences in the multicopy ribosomal internal transcribed spacer (ITS) region suggested the possible hybrid origin of two European soft coral species, Alcyonium hibernicum and Bellonella bocagei. To examine this possibility further we cloned and sequenced ITS-1 from multiple individuals and populations of these two species as well as two sympatric congeners, A. coralloides and A. sp. M 2. Phylogenetic analyses separated the observed sequence variants into two distinct clades. All A. coralloides sequences belonged to clade A, while A. sp. M 2 had only clade B sequences. A majority of A. hibernicum individuals, however, contained both clade A and B sequences that were identical to the predominant sequence variants found in A. coralloides and A. sp. M 2, respectively. This pattern of additivity suggests that A. hibernicum originated from a hybrid cross between A. coralloides and A. sp. M 2, a hypothesis that is supported by its unusual mode of reproduction (meiotic parthenogenesis). The predominant sequence variant found in B. bocagei was a unique, derived clade B sequence; in addition, however, most individuals of this species also had copies of a sequence identified as a recombinant between clade A and clade B sequence types. The presence of this recombinant sequence in the B. bocagei genome suggests that this species may also be the product of past hybridization events within the clade. Reticulate evolution may explain the failure of several previous studies to resolve the phylogeny of these four species.

Cnidarians are early-diverging metazoans, but evolutionary aspects of some taxa are still poorly understood, as in the order Zoantharia (Anthozoa: Hexacorallia). Zoantharians have been divided into two suborders based on the arrangement of the fifth septae as complete (Macrocnemina) or incomplete (Brachycnemina). Previous Data S1. Information on the mitogenome sizes and gene order arrangements for each of the specimens investigated. Data S2. Pairwise genetic distances of 13 mitochondrial protein-coding genes. Alignment S1. Mitogenome coding DNA sequence alignment. Alignment S2. 16S alignment. Alignment S3. Atp6 alignment. Alignment S4. Atp8 alignment. Alignment S5. Cox1 alignment. Alignment S6. Cox2 alignment. Alignment S7. Cox3 alignment. Alignment S8. Cytb alignment. Alignment S9. Nad1 alignment. Alignment S10. Nad2 alignment. Alignment S11. Nad3 alignment. Alignment S12. Nad4 alignment. Alignment S13. Nad4l alignment. Alignment S14. Nad5 alignment. Alignment S15. Nad6 alignment. How to cite this article: Poliseno A, Santos MEA, Kise H, et al. Evolutionary implications of analyses of complete mitochondrial genomes across order Zoantharia (Cnidaria: Hexacorallia).

Several recent studies have suggested that hybridization may play a previously unrecognized and important role in the evolution of corals. Our observations of polymorphic and recombinant sequences in the multicopy ribosomal internal transcribed spacer (ITS) region suggested the possible hybrid origin of two European soft coral species, Alcyonium hibernicum and Bellonella bocagei. To examine this possibility further we cloned and sequenced ITS-1 from multiple individuals and populations of these two species as well as two sympatric congeners, A. coralloides and A. sp. M 2. Phylogenetic analyses separated the observed sequence variants into two distinct clades. All A. coralloides sequences belonged to clade A, while A. sp. M 2 had only clade B sequences. A majority of A. hibernicum individuals, however, contained both clade A and B sequences that were identical to the predominant sequence variants found in A. coralloides and A. sp. M 2, respectively. This pattern of additivity suggests that A. hibernicum originated from a hybrid cross between A. coralloides and A. sp. M 2, a hypothesis that is supported by its unusual mode of reproduction (meiotic parthenogenesis). The predominant sequence variant found in B. bocagei was a unique, derived clade B sequence; in addition, however, most individuals of this species also had copies of a sequence identified as a recombinant between clade A and clade B sequence types. The presence of this recombinant sequence in the B. bocagei genome suggests that this species may also be the product of past hybridization events within the clade. Reticulate evolution may explain the failure of several previous studies to resolve the phylogeny of these four species.

Background: Curious parallels between the processes of species and language evolution have been observed by many researchers. Retracing the evolution of Indo-European (IE) languages remains one of the most intriguing intellectual challenges in historical linguistics. Most of the IE language studies use the traditional phylogenetic tree model to represent the evolution of natural languages, thus not taking into account reticulate evolutionary events, such as language hybridization and word borrowing which can be associated with species hybridization and horizontal gene transfer, respectively. More recently, implicit evolutionary networks, such as split graphs and minimal lateral networks, have been used to account for reticulate evolution in linguistics. Results: Striking parallels existing between the evolution of species and natural languages allowed us to apply three computational biology methods for reconstruction of phylogenetic networks to model the evolution of IE languages. We show how the transfer of methods between the two disciplines can be achieved, making necessary methodological adaptations. Considering basic vocabulary data from the well-known Dyen's lexical database, which contains word forms in 84 IE languages for the meanings of a 200-meaning Swadesh list, we adapt a recently developed computational biology algorithm for building explicit hybridization networks to study the evolution of IE languages and compare our findings to the results provided by the split graph and galled network methods. Conclusion: We conclude that explicit phylogenetic networks can be successfully used to identify donors and recipients of lexical material as well as the degree of influence of each donor language on the corresponding recipient languages. We show that our algorithm is well suited to detect reticulate relationships among languages, and present some historical and linguistic justification for the results obtained. Our findings could be further refined if relevant syntactic, phonological and morphological data could be analyzed along with the available lexical data.

Cladistic analyses of the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) sequences from 55 samples corresponding to 34 taxa in the genus Armeria reveal that ITS sequence diversity among and within species utterly conflicts with patterns of morphological similarity. Three facts are apparent from the results here reported: (1) different samples of a single subspecies, A. villosa subsp. longiaristata, appear in three of the five major clades; (2) samples of at least one of the six subspecies of A. villosa appear in four of the five major clades; and (3) the composition of major clades shows greater congruence with the geographic origin of plants than with the traditional systematic arrangement based primarily on morphology. Specifically, the clades here termed Ia, II, III, and IV each encompass terminals restricted to geographically delimited areas. There are alternative explanations for the ITS pattern, but the most likely one is that nucleotide positions supporting the major clades are due, in some of the samples, to concerted evolution following horizontal transfer (gene flow) rather than to recency of common ancestry. This interpretation is consistent with previous systematic and experimental evidence and implies that reticulation in Armeria may be extensive. Harlan and de Wet (1963, Evolution 17:497-501) proposed the compilospecies concept to account for situations in which a genetically "aggressive" species captures portions of the genome of other sympatric species by means of extensive introgression. Evidence of extensive reticulation , ecological diversification, and geographic pattern indicates that A. villosa may fit the compilospecies concept, which is here supported on molecular grounds for the first time.

Species of Armeria from the southern part of the Iberian Peninsula have been postulated to undergo extensive reticulate evolution based on ITS sequences as well as morphometric, ecogeographical evidence and crossing experiments. The hypothesis that a rare serpentine endemic (A. villosa subsp. carratracensis) originated from a cross between another serpentine tolerant endemic (A. colorata) and a widespread limestone species (A. villosa subsp. longiaristata) is tested with molecular markers. Sequences from two chloroplast regions [trnL (UAA)-trnF (GAA) and trnD (GUC)-trnT (GGU)], from nuclear ribosomal DNA (ITS-1 + 5.8S + ITS-2) as well as RAPD data are used. Variation in chloroplast and nuclear sequence data shows a clear geographical structure irrespective of taxonomic ascription. The putative hybrid taxon shares locally restricted haplotypes and ribotypes both with A. colorata and populations of A. villosa subsp. longiaristata. Species speci®c markers for the progenitors have not been found and thus a divergent scenario cannot be conclusively excluded. A principal coordinate analysis as well as a UPGMA analysis of the RAPD data based on the Dice coecient are consistent with the hybrid origin. A joint examination of the various markers together with the previous biological evidence and the geographical structure of the chloroplast and ITS markers raises the likelihood of the reticulate hypothesis. Reasons for inconclusiveness are discussed. Using PCR-RFLPs of arti-®cially synthesized hybrids and backcrosses, evidence for a maternal inheritance of chloroplast DNA in Armeria is provided.

The Malva alliance is a well-defined group with extensive morphological homoplasy. As a result, the relationships among the taxa as well as the evolution of morphological traits have remained elusive and the traditional classifications are highly artificial. Using five molecular markers (nuclear ITS, plastid matK plus trnK, ndhF, trnL-trnF, psbA-trnH), we arrived at a phylogenetic hypothesis of this group, the genera Alcea, Althaea and Malvalthaea being studied here for the first time with molecular data. Althaea and, in particular, Lavatera and Malva are highly polyphyletic as currently circumscribed, because their diagnostic characters, the number and degree of fusion of the epicalyx bracts, evolve in a highly homoplasious manner. In contrast, fruit morphology largely agrees with the molecularly delimited groups. Hybrid origins confirmed for the genus Malvalthaea and for Lavatera mauritanica and hybridization in the group of ruderal small-flowered mallows underline the importance of reticulate evolution in shaping the history of this group and complicating the interpretation of morphological evolution.

A parsimony analysis of 133 sequences of the nuclear ribosomal DNA ITS1 + 5.8S + ITS2 region from 71 taxa in Armeria was carried out. The presence of additive polymorphic sites (APS; occurring in 14 accessions) fits the reticulate scenario proposed in previous work for explaining the ITS pattern of variation on a much smaller scale and is based mainly on the geographical structure of the data, irrespective of taxonomic boundaries. Despite the relatively low bootstrap values and large polytomies, part of which are likely due to disruptive effects of reticulation and concerted evolution in these multicopy sequences, the ITS analysis has phylogenetic and biogeographic implications. APS detected in this study are consistent with hypothesized hybridization events, although biased concerted evolution, previously documented in the genus, needs to be invoked for specific cases and may be responsible for a possible ''sink'' effect in terminals from a large clade. The causes for sequences of the same species appearing in different clades (here termed transclade) are discussed.

Background Sea-lavenders (Limonium Mill., Plumbaginaceae) are a cosmopolitan group of diploid and polyploid plants often adapted to extreme saline environments, with a mostly Tethyan distribution, occurring in the Mediterranean, Irano-Turanian, Euro-Siberian and in the New World. The halophylic Limonium vulgare polyploid complex in particular, presents a large distribution throughout extreme salt-marsh habitats and shows little morphological but high taximetric variation, frequently blurring species delimitation. In this work we pursue three main goals: assert whether SNP data from polyploid individuals has the resolution to distinguish the seven sampled species, to better understand how genetically structured Limonium vulgare is, and attempt to identify specific molecular mechanisms for the differentiation between L. maritimum and L. vulgare. For this purpose, 95 individuals were genotyped using Genotyping by Sequencing (GBS), which were assembled as two independent datasets using ...

Limonium vulgare and related species form a complex group, but until now cytological and genetic studies have been based on single species and specific geographical areas. We investigated genome size, karyological and genetic diversity in samples from Western Mediterranean and evaluated the phylogenetic relationships among the species of this complex. Genome size was assessed using flow cytometry on samples from natural populations of L. vulgare, L. maritimum and L. narbonense. Chromosome counts were conducted in plants obtained from seeds collected in the field. The internal transcribed spacer ITS1 of the nuclear rDNAs was used to assess ITS polymorphisms as well as the phylogenetic relationships within the L. vulgare complex. Our analyses showed that all species were tetraploid, with the chromosome number of L. maritimum being presented here for the first time. Significant differences were observed in genome size, with L. narbonense having lower genome sizes than the other two species, and possible aneuploids being detected. Ten new ITS sequences from L. vulgare, L. narbonense and L. maritimum were provided. Most species' populations showed unique ribotypes, and L. narbonense has the highest ribotype diversity. One of the L. maritimum populations presented a closer genetic relationship with L. vulgare, whereas the other two seemed to be more related with L. narbonense. Phylogenetic analyses confirmed that L. vulgare and L. narbonense form a monophyletic group, sister to the remaining Limonium species. Our results put into evidence that the studied species may represent a relatively early stage of divergence.

The genus Limonium (sea lavenders) is characterized by various taxonomically challenging complexes with great karyological diversity, and chromosomes ranging in size from diminutive to large. In this biosystematic study, our goal was to investigate cytogenetic features of twelve sexual and asexual taxa in a particular part of their distribution range. Genome size, chromosome number variation, and other cytological features were investigated in a representative set of sea lavenders with distinct leaf venation, present in the coastal areas of the Iberian Peninsula, Morocco, France, and the Channel Islands. Flow cytometric genome size determinations were made in plants from natural populations. Cell and nuclear areas of epidermal cells, as well as, chromosome numbers and length were measured in plants from ex situ collections. Our findings revealed positive and significant correlations between holoploid genome size and cell size, nuclei area and chromosome total length. Tetraploid taxa...