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Resolution of deep nodes yields an improved backbone phylogeny and a new basal lineage to study early evolution of Asteraceae
Abstract
A backbone phylogeny that fully resolves all subfamily and deeper nodes of Asteraceae was constructed using 14 chloroplast DNA loci. The recently named genus Famatinanthus was found to be sister to the Mutisioideae-Asteroideae clade that represents more than 99% of Asteraceae and was found to have the two chloroplast inversions present in all Asteraceae except the nine genera of Barnadesioideae. A monotypic subfamily Famatinanthoideae and tribe Famatinantheae are named herein as new. Relationships among the basal lineages of the family were resolved with strong support in the Bayesian analysis as (Barnadesioideae (Famatinanthoideae (Mutisioideae (Stifftioideae (Wunderlichioideae-Asteroideae))))). Ancestral state reconstruction of ten morphological characters at the root node of the Asteraceae showed that the ancestral sunflower would have had a woody habit, alternate leaves, solitary capitulescences, epaleate receptacles, smooth styles, smooth to microechinate pollen surface sculpturing, white to yellow corollas, and insect-mediated pollination. Herbaceous habit, echinate pollen surface, pubescent styles, and cymose capitulescences were reconstructed for backbone nodes of the phylogeny corresponding to clades that evolved shortly after Asteraceae dispersed out of South America. No support was found for discoid capitula, multiseriate involucres or bird pollination as the ancestral character condition for any node. Using this more resolved phylogenetic tree, the recently described Raiguenrayun cura + Mutisiapollis telleriae fossil should be associated to a more derived node than previously suggested when time calibrating phylogenies of Asteraceae.
... The family Asteraceae is the second-largest flowering plant family after the Orchidaceae (Umber et al., 2022). It presents a wide geographical distribution occupying almost all the habitats and vegetation formations, although it is predominant in tropical and subtropical regions, mainly in open areas, such as grasslands (Anderberg et al., 2007;Çitak et al., 2021), with around 13 subfamilies, 44 tribes, over 1600 genera, and 23,000 species (Funk et al., 2009;Panero et al., 2014). The family Asteraceae is included in the Asterids clade of the order Asterales according to APG III (2009). ...
... This classification is used in some floras and handbooks e.g., in the Flora Reipublicae Popularis Sinicae (Ling et al., 1985). However, there have been major changes in the classification of Asteraceae in recent decades with a better phylogenetic framework (Jansen and Palmer, 1987;Kim and Jansen, 1995;Bayer and Starr, 1998;Kim et al., 2002;Panero andFunk, 2002, 2008;Goertzen et al., 2003;Panero, 2005;Funk et al., 2009;Smith et al., 2009;Torices, 2010;Panero et al., 2014;Mandel et al., 2015). Based on the 10 or 14 chloroplast DNA (cpDNA) markers, Panero andFunk (2002, 2008) and Panero et al. (2014) reconstructed the backbone of Asteraceae with 12-13 major clades (subfamilies) identified. ...
... However, there have been major changes in the classification of Asteraceae in recent decades with a better phylogenetic framework (Jansen and Palmer, 1987;Kim and Jansen, 1995;Bayer and Starr, 1998;Kim et al., 2002;Panero andFunk, 2002, 2008;Goertzen et al., 2003;Panero, 2005;Funk et al., 2009;Smith et al., 2009;Torices, 2010;Panero et al., 2014;Mandel et al., 2015). Based on the 10 or 14 chloroplast DNA (cpDNA) markers, Panero andFunk (2002, 2008) and Panero et al. (2014) reconstructed the backbone of Asteraceae with 12-13 major clades (subfamilies) identified. The Chinese Asteraceae comprised approximately 2336 indigenous (1145 endemic) and 248 genera nearly 15% of world genera (Shih et al., 2011) belonging to 22 tribes and 7 subfamilies (Fu et al., 2016). ...
The Asteraceae is one of the most taxonomically complex, medicinally and economically important family among the angiosperms. Therefore, this study aims to evaluate the leaf anatomical features to correctly identify the Asteraceae species of Hainan Island. Scanning electron microscopy was used to investigate various micro epidermal traits. Leaf micromorphology of all the species was characterized, visualized and compared using UPGMA Cluster analysis (CA), chord diagram and ridgeline plot. Generally, the trichomes were divided into two main categories, glandular trichomes (GTs) and non-glandular trichomes (NGTs). The GTs was further classified into capitate, subsessile and sessile capitate. The NGTs were classified into long falcate, short conical, lipidote and filiform. The stomata were mostly anomocytic and anisocytic sometimes mixed with tetracytic type. However, there are varied assortments of trichomes and stomata with transitions among them especially in their type, frequency and number of cells. The trichomes and stomata can be successfully used for the delimitation of the 2 genera within the family. The statistical analysis can help in better understanding of the results that characterized, visualized and compared the leaf anatomical traits of the Asteraceae taxa. Overall, these traits collectively have a significant taxonomic potential to identify and define species boundaries at the tribe and generic levels and can be used as an additional tool for regrouping taxa within Asteraceae.
... Morphological and molecular evolutionary analyses require a well-resolved phylogeny. Previous analyses, largely using plastid genes, have supported the monophyly of Asteraceae (Anderberg et al., 2007), defining 45 tribes in 13 subfamilies Panero et al., 2014). The relationships among the Asteraceae subfamilies and most tribes have also been resolved using plastid sequences (Fu et al., 2016;Panero and Crozier, 2016), although some uncertainties remain. ...
... Asteraceae nuclear phylogeny reveals highly supported relationships among subfamilies To reconstruct a tribal-level Asteraceae phylogeny, 243 Asteraceae species were sampled, representing all 13 subfamilies and 41 of the 45 recognized tribes (Panero and Funk, 2008;Funk et al., 2009b;Panero et al., 2014;Fu et al., 2016;Huang et al., 2016b) (149 species in Asteroideae, e.g., sunflowers, daisies, and chrysanthemums; 27 in Cichorioideae, e.g., lettuce and dandelion; 33 in Carduoideae, e.g., artichoke and thistles; and five outgroup taxa). We newly generated transcriptome and genome sequences from 121 (for one species, RNAs from two samples were sequenced) and 16 species, respectively (see Supporting Information; Tables S1, S2), plus 67 previous datasets from our lab (Zeng et al., 2014;Liu et al., 2015;Huang et al., 2016b) and 44 publicly available datasets (Tables S1, S2). ...
... Phylogenetic analyses using both coalescent and maximum likelihood (ML) methods with multiple datasets comprising nuclear genes yielded highly supported and consistent Asteraceae phylogenies (Figures 1, 2, S3, S14) (see Supporting Information for details). Asteraceae were monophyletic in all analyses here, forming a sister clade to Calyceraceae (three genera sampled), and the phylogeny was mostly consistent with previously reported topologies (e.g., Panero and Funk, 2008;Panero et al., 2014;Fu et al., 2016;Panero and Crozier, 2016). Asteroideae and seven other subfamilies were monophyletic with 100% support in all analyses; Famatinanthoideae and Hecastocleidoideae were monotypic; whereas Wunderlichioideae, Cichorioideae, and Carduoideae were not monophyletic ( Figure 1). ...
Biodiversity is not evenly distributed among related groups, raising questions about the factors contributing to such disparities. The sunflower family (Asteraceae, >26,000 species) is among the largest and most diverse plant families, but its species diversity is concentrated in a few subfamilies, providing an opportunity to study the factors affecting biodiversity. Phylotranscriptomic analyses here of 244 transcriptomes and genomes produced a phylogeny with strong support for the monophyly of Asteraceae and the monophyly of most subfamilies and tribes. This phylogeny provides a reference for detecting changes in diversification rates and possible factors affecting Asteraceae diversity, which include global climate shifts, whole‐genome duplications (WGDs), and morphological evolution. The origin of Asteraceae was estimated at ~83 Mya, with most subfamilies having diverged before the Cretaceous–Paleocene boundary. Phylotranscriptomic analyses supported the existence of 41 WGDs in Asteraceae. Changes to herbaceousness and capitulescence with multiple flower‐like capitula, often with distinct florets and scaly pappus/receptacular bracts, are associated with multiple upshifts in diversification rate. WGDs might have contributed to the survival of early Asteraceae by providing new genetic materials to support morphological transitions. The resulting competitive advantage for adapting to different niches would have increased biodiversity in Asteraceae.
... Processes as whole-genome duplication and hybridization are common in Asteraceae (Ellstrand et al., 1996;Barker et al., 2008;Symonds et al., 2010;Soto-Trejo et al., 2013;Huang et al., 2016) and can obscure evolutionary patterns and hinder the recognition and delimitation of clades. Currently, 12-13 subfamilies and between 35 and 45 tribes are recognized (Anderberg et al., 2007;Funk et al., 2009;Panero et al., 2014;Fu et al., 2016;Panero & Crozier, 2016). The efforts to resolve relationships within the family have resulted in robust phylogenetic hypothesis (Funk et al., 2009;Panero et al., 2014;Huang et al., 2016;Panero & Crozier, 2016;Mandel et al., 2017Mandel et al., , 2019. ...
... Currently, 12-13 subfamilies and between 35 and 45 tribes are recognized (Anderberg et al., 2007;Funk et al., 2009;Panero et al., 2014;Fu et al., 2016;Panero & Crozier, 2016). The efforts to resolve relationships within the family have resulted in robust phylogenetic hypothesis (Funk et al., 2009;Panero et al., 2014;Huang et al., 2016;Panero & Crozier, 2016;Mandel et al., 2017Mandel et al., , 2019. The incorporation of nuclear and chloroplast data, obtained from next-generation sequencing (NSG) methods, has allowed resolving problematic branches and increased the support of clades (Huang et al., 2016;Mandel et al., 2014Mandel et al., , 2015Mandel et al., , 2017Mandel et al., , 2019. ...
... North America, particularly Mexico, represents an important site for the diversification of some of the most diverse lineages in the family, such as the "Heliantheae alliance" clade within the subfamily Asteroideae (Noyes & Rieseberg, 1999;Suárez-Mota & Villaseñor, 2011;Villaseñor, 2018). Evidence suggests that this clade is the result of a single colonization event from Africa or Asia to North America 22-35 million years ago (Panero, 2007;Funk et al., 2009;Panero et al., 2014;Panero & Crozier, 2016). The Heliantheae alliance is composed of 13 tribes, including Athroismeae, Bahieae, Chaenactideae, Coreopsideae, Eupatorieae, Helenieae, Heliantheae, Madieae, Millerieae, Neurolaeneae, Perityleae, Polymnieae, and Tageteae (Anderberg et al., 2007). ...
Asteraceae is the largest plant family in México with about 417 genera and 3,113 species, more than 60% of them endemic. Phylogenetic relationships at subfamily and tribal levels have been previously resolved employing both nuclear and plastid molecular markers. However, Asteraceae species native to Mexico have been underrepresented in such phylogenies. To tackle this issue, the taxon sampling of this study included 90 Asteraceae species native to México, four species from the Caribbean, 119 previously sequenced species, and six outgroups. With this sampling, all the Asteraceae subfamilies and all of the tribes recognized to date are represented. The analyzed data set consisted of eleven chloroplast markers (atpB, matK, ndhC, ndhD, ndhF, ndhI, ndhJ, ndhK, rbcL, trnL‐trnF, 23S‐trnA). We present two phylogenetic reconstructions obtained by maximum likelihood and pseudocoalescent methods. Besides, we present a time‐calibrated phylogeny, used to infer the best configuration of diversification rate shifts. Our results show that Mexican species are distributed mainly in the subfamily Asteroideae (80 species), followed by Cichorioideae (6 species), Carduoideae (2 species), and Mutisioideae (2 species). Four net diversification rate shifts were found: One near the base of the tree and four within Asteroideae subfamily. Our extended sampling of the family with the representation of native species to Mexico allowed us to identify important events in the evolutionary history of the family.
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... Over time, various classification systems have emerged, with two notable ones being proposed by Cassini (1816) comprising 19 tribes, and Bentham (1873) introducing two subfamilies (Asteroideae and Cichorioideae) along with 13 tribes. Subsequent revisions have expanded these classifications, evolving into 13 subfamilies (major clades) and 43 tribes in more recent systems (Panero and Funk 2002;Funk et al. 2009;Panero et al. 2014). Due to the vast number of species and the global geographic distribution of Asteraceae, undertaking taxonomic and evolutionary investigations within this plant family poses a formidable challenge (Mandel et al. 2019). ...
... In our cluster microechinate types represent the basal tribes which were first to separate and clearly supporting the trend of cluster presented by Teller� ıa et al. (2023). Our clustering approach aligns with the classification system of Asteraceae (Funk et al. 2009;Panero et al. 2014), which groups species within the same subfamily closely together in the cluster. This alignment underscores the importance of palynotaxonomy in conjunction with other morphological features. ...
... Chen and Anderberg (2011) even considered it to be a genus of uncertain placement (genera Incertae sedis) in Flora of China. More recently, studies have sought to elucidate the phylogenetic position and relationship of Cavea (Panero and Funk 2008;Anderberg and Ohlson 2012;Panero et al. 2014;Mandel et al. 2019). Based on limited sequences, the present molecular studies supported the close relationship between two monotypic genera, Cavea and Gymnarrhena Desf. ...
... The megafamily Asteraceae is composed of 13 subfamilies (backbone of the Asteraceae, Barnadesioideae, Famatinanthoideae, Mutisioideae, Gochnatioideaet, Stifftioideae, Wunderlichioideae, Hecastocleidoideae, Carduoideae, Pertyoideae, Gymnarrhenoideae, Cichorioideae, Corymbiodeae, and Asteroideae (Panero and Funk 2008;Panero et al. 2014;Mandel et al. 2019). With the fast development of molecular sequencing technology, to date, about 754 Asteraceae plastomes samples have been sequenced, but only seven subfamilies (Asteroideae, Barnadesioideae, Carduoideae, Cichorioideae, Pertyoideae, Mutisioideae and Wunderlichioideae) are covered (1 June 2021, the National Center for Biotechnology Information, NCBI, https://www.ncbi.nlm. ...
The family Asteraceae includes about 10% of angiosperm species. The tribe Gymnarrheneae is an excellent example of a nonmissing link tribe. It contains two monotypic genera (Cavea and Gymnarrhena) and is positioned phylogenetically to a large clade of Asteroideae ? Corymbiodeae ? Cichorioideae which contain more than 80% of all the Asteraceae. The genus Cavea (tribe Gymnar-rheneae) is a medicinally significant genus and relatively rare in the Qinghai-Tibet Plateau (QTP) and its adjacent areas. Based on the limited evidence, resolving phylogenetic relationship of the genus Cavea has proven difficult for a long time. Despite its important medicinal value, genomic resources of Cavea are still lacking, preventing our understanding of its evolutionary history. In recent years, the plastid genomes are widely used in phylogenetic analysis. To provide more useful genetic data for resolving the systematic disputation of Cavea, here, a complete chloroplast genome of Cavea tanguensis was obtained using Illumina sequencing data for the first time. The whole circular cp genome of C. tanguensis was 150,799 bp in length, contained a large single-copy (LSC) region of 82,514 bp and a small single-copy (SSC) region of 18,471 bp. These two regions were separated by a pair of inverted repeat regions (IRa and IRb), each of them being 24,907 bp in length. A total of 135 functional genes were annotated which consisted of 89 protein-coding genes, 38 trnA genes, and eight rRNA genes. The overall GC content of the chloroplast genome sequence was 37.5%, and the GC contents of the LSC, SSC and IR regions were 35.6%, 30.9% and 37.0%, respectively. We analysed insertions/deletions, and simple-sequence repeats in the chloroplast genomes, and discovered relatively highly variable regions (trnD-GUC/rpoB, trnL-UCC/ndhK, and ycf 1) that will potentially provide plastid markers for further taxonomic, phylogenetic, and population genetic studies in Asteraceae. The phylogenetic analyses based on 22 complete chloroplast genome sequences robustly supported that C. tanguensis formed a sister group with the subfamilies Asteroideae and Car-duoideae, consistent with the finding of recent studies. This study provides new insight into the plastid genome evolution and phylogenetic relationships. Moreover, it would be fundamental to formulate potential conservation and management strategies for the enigmatic species in the Himalaya.
... Species of the family Asteraceae exist in every type of habitat and every continent including Antarctica (Barreda et al., 2015;Mandel et al., 2019). This family is divided into 13 subfamilies (Mandel et al., 2019;Panero & Crozier, 2016;Panero et al., 2014). ...
... Watson et al. (2020) referred to the five tribes Senecioneae, Astereae, Anthemideae, Gnaphalieae, and Calenduleae as the Fab(ulous) Five. They are taxonomically difficult tribes, and conflicting phylogenetic signals were recorded for these tribes based on the plastid, nuclear, and transcriptomic data (for details see the discussion) (Fu et al., 2016;Mandel et al., 2019;Panero & Crozier, 2016;Panero et al., 2014;Watson et al., 2020). ...
The genus Blumea (Asteroideae, Asteraceae) comprises about 100 species, including herbs, shrubs, and small trees. Previous studies have been unable to resolve taxonomic issues and the phylogeny of the genus Blumea due to the low polymorphism of molecular markers. Therefore, suitable polymorphic regions need to be identified. Here, we de novo assembled plastomes of the three Blumea species B. oxyodonta, B. tenella, and B. balsamifera and compared them with 26 other species of Asteroideae after correction of annotations. These species have quadripartite plastomes with similar gene content, genome organization, and inverted repeat contraction and expansion comprising 113 genes, including 80 protein‐coding, 29 transfer RNA, and 4 ribosomal RNA genes. The comparative analysis of codon usage, amino acid frequency, microsatellite repeats, oligonucleotide repeats, and transition and transversion substitutions has revealed high resemblance among the newly assembled species of Blumea. We identified 10 highly polymorphic regions with nucleotide diversity above 0.02, including rps16‐trnQ, ycf1, ndhF‐rpl32, petN‐psbM, and rpl32‐trnL, and they may be suitable for the development of robust, authentic, and cost‐effective markers for barcoding and inference of the phylogeny of the genus Blumea. Among these highly polymorphic regions, five regions also co‐occurred with oligonucleotide repeats and support use of repeats as a proxy for the identification of polymorphic loci. The phylogenetic analysis revealed a close relationship between Blumea and Pluchea within the tribe Inuleae. At tribe level, our phylogeny supports a sister relationship between Astereae and Anthemideae rooted as Gnaphalieae, Calenduleae, and Senecioneae. These results are contradictory to recent studies which reported a sister relationship between “Senecioneae and Anthemideae” and “Astereae and Gnaphalieae” or a sister relationship between Astereae and Gnaphalieae rooted as Calenduleae, Anthemideae, and then Senecioneae using nuclear genome sequences. The conflicting phylogenetic signals observed at the tribal level between plastid and nuclear genome data require further investigation.
... The tribe Astereaehas has~222 genera and~3,100 species, which is the second largest tribe of Asteraceae (Noyes and Rieseberg, 1999;Brouillet et al., 2004;Panero et al., 2004;Panero and Crozier, 2016;Fu et al., 2016). The tribe Senecioneae has over 150 genera and 3,500 species (Nordenstam, 2007), more than the species number of the tribe Astereae. ...
Aster L. is an economically and phylogenetically important genus in the tribe Astereae. Here, the complete plastomes of the eight Aster species were assembled and characterized using next-generation sequencing datasets. The results indicated the complete plastomes of Aster had a quadripartite structure. These genomes were 152,045–152,729 bp in length and contained 132–133 genes, including 87 protein-coding genes, 37–38 tRNA genes, and eight rRNA genes. Expansion or contraction of inverted repeat regions and forward, palindromic, complement, and reverse repeats were detected in the eight Aster species. Additionally, our analyses showed the richest type of simple sequence repeats was A/T mononucleotides, and 14 highly variable regions were discovered by analyzing the border regions, sequence divergence, and hotspots. Phylogenetic analyses indicated that 27 species in Astereae were clustered into six clades, i.e., A to D, North American, and outgroup clades, and supported that the genera Heteropappus, Kalimeris, and Heteroplexis are nested within Aster. The results indicated the clades B to D might be considered as genera. Divergence time estimate showed the clades A, B, C, and D diverged at 23.15 Mya, 15.13 Mya, 24.29 Mya, and 21.66 Mya, respectively. These results shed light on the phylogenetic relationships of Aster and provided new information on species identification of Aster and its related genera.
... The species Chrysanthemum indicum L. and Dendranthema indicum (L.) Des Moul. were selected as the outgroup following previous works Funk, 2002, 2008;Panero et al., 2014). ...
Aster yaoshanensis sp. nov., a new species of the family Asteraceae is here described and illustrated. The species is presently known only from rock crevices of mountain valleys in Dayaoshan National Nature Reserve, Guangxi autonomous region, China. Phylogenetic analyses based on ITS sequences and complete plastome data have shown that this new species is a member of genus Aster with high support. Morphologically, it mostly resembles A. jishouensis, but it can be easily distinguished from the latter by bract indumentum (glabrous except margin ciliate vs. villous especially on veins abaxially, glabrous adaxially) and color (green vs. purple), shorter corolla (3.2–3.5 mm vs. 4.5–5.3 mm), bract stalk (obvious, ca.1.2 mm vs. sessile), and different distribution (Guangxi vs. Hunan). The detailed description, distribution map, and photos are provided. This study further elucidates the species identification, phylogeny and characteristic evolution of Aster.
... This knowledge was reflected in the number of species (n = 185) and families mentioned (n = 57) that are used in their crops. The plant families most mentioned by farmers (Asteraceae, Lamiaceae, and Brassicaceae) were also quite prominent in other ethnobotanical studies undertaken in rural communities, quilombolas, and Indigenous areas (Ávila et Asteraceae is one of the largest botanical families, comprising approximately 1,700 genera and 27,000 species; it comprises 10% of all angiosperm species (Funk et al. 2009;Panero et al. 2014;Panero & Crozier 2016). There are approximately 289 genera and 2,097 species of Asteraceae in Brazil, distributed in essentially all of its vegetation formations (Flora do Brasil 2020, continuously updated). ...
Rural communities have repertoires of knowledge associated with the use of plants related to various social and biological aspects. The objective of this study was to identify the diversity of use of food and medicinal plants by the Brejal rural community in Rio de Janeiro state, Brazil. Ethnobotanical data was obtained using the snowball technique and semi-structured interviews with 22 farmers. Species diversity and use-value indices were calculated. Relationships among the numbers of plants cited, interviewee age, gender, and working time in agriculture were investigated. We identified a high diversity index, comprising 185 species belonging to 53 botanical families, notably Asteraceae, Lamiaceae, and Brassicaceae. The greatest use-values were associated with exotic species. Leaves were the plant structures most utilized, and infusions the predominant method of preparation. The prescriptions most commonly cited related to illnesses of the digestive system. Men and women did not differ in terms of the numbers of species cited. We found no correlations between the ages of the interviewees and their working time in agriculture. The uniformity of the number of citations demonstrates the shared knowledge throughout the community, covering a high diversity of plants, uses, indications and preparations.
... At the same time, because early studies had lower levels of resolution and obtaining complete genomes was di cult, small and easily obtained chloroplast genes became more prominent. For early evolutionary studies in Asteraceae, markers from the chloroplast genome were used [29,30,31] . Compared with constructing phylogenetic trees using a single gene or a few markers, some families have used the complete chloroplast genome CDS gene set to reconstruct their phylogenetic relationships. ...
Dahlia pinnata Cavanilles 1791 is an important ornamental plant worldwide. The chloroplast genome has obvious advantages in studies of systematic evolution at the plant classification and species level, making it an important resource for phylogenetic research.Here, we sequenced the full chloroplast genome from D. pinnata 'Chocolate' and found that it exhibited a typical tetrad structure. The full-length D. pinnata chloroplast genome was 152,107 bp, with a GC content of 38.45%. The genome included an 83,704 bp large single-copy (LSC) region, an 18,347 bp small single-copy (SSC) region, and a pair of 25,028 bp inverted repeats ( IRa and IRb ). A total of 134 genes were annotated, including 86 protein-coding genes, 38 transfer RNA genes, 8 ribosomal RNA genes, and 2 pseudogenes. Analysis of password preferences shows that passwords chosen by D. pinnata tend to end with A/U. A total of 161 SSR markers were detected in the simple sequence repeat (SSR) analysis. Phylogenetic analysis that the Dahlia species formed a monophyly. Dahlia was clustered with Cosmos - Bidens , which differed from studies using nuclear genomic DNA. We suggest that nuclear-cytoplasmic incongruences may be widespread in Asteraceae, and should be thoroughly evaluated in order to understand the true evolutionary history of this economically-important group of plants.
... Various kinds of studies have been conducted in an attempt to help explain the diversity of species, habitats, life forms and life cycles of the Asteraceae. The ideas/studies about the reasons for the high diversification rates of Asteraceae include whole-genome duplication (polyploidization) (WGD; Zhang et al., 2021a), genetic diversity (Pascual-Díaz et al., 2021), interaction with insects that serve as pollinators (Panero et al., 2014) and production of secondary metabolites that deter predators especially insects (Seaman, 1982;Vanderplanck et al., 2020). An inflorescence compressed into a head or capitulum is viewed as a pseudanthia or false flower (Zhang and Elomaa, 2021). ...
The Asteraceae with up to 30,000 species occurs on all continents except Antarctica and in all major vegetation zones on earth. Our primary aim was to consider cypselae dormancy-break and germination of Asteraceae in relation to ecology, vegetation zones and evolution. Cypselae are desiccation-tolerant and in various tribes, genera, species and life forms of Asteraceae are either non-dormant (ND) or have non-deep physiological dormancy (PD) at maturity. All six types of non-deep PD are found among the Asteraceae, and dormancy is broken by cold or warm stratification or by afterripening. Soil cypselae banks may be formed but mostly are short-lived. Much within-species variation in dormancy-break and germination has been found. Using data compiled for 1192 species in 373 genera and 35 tribes of Asteraceae, we considered ND and PD in relation to life form, vegetation zone and tribe. Senecioneae and Astereae had the best representation across the vegetation zones on earth. In evergreen and semi-evergreen rainforests, more species have ND than PD, but in all other vegetation zones, except alpine/high-latitude tundra (where ND and PD are equal), more species have PD than ND. Tribes in the basal and central grades and those in the Heliantheae Alliance have both ND and PD. The high diversity and lability of non-deep PD may have enhanced the rate of species diversification by promoting the survival of new species and/or species in new habitats that became available following globally disruptive events since the origin of the Asteraceae in the Late Cretaceous.
... Genome-scale data including plastid and nuclear genome data have significantly increased the number of informative characters for phylogenetic analyses and have been successfully utilized for reconstructing phylogenies of many plant groups recently Xiang et al., 2017;Kleinkopf et al., 2019;Leebens-Mack et al., 2019;Li et al., 2019;Medina et al., 2019;Ma et al., 2021). The chloroplast genome sequences, easily obtainable from genome skimming (Jansen & Ruhlman, 2012), have become a popular source of data for reconstructing the phylogenetic relationships of many groups at different levels in the past ten years (Bock et al., 2014;Panero et al., 2014;Zhang et al., 2015Zhang et al., , 2016Wen et al., 2018;Liu et al., 2019;Valcárcel & Wen, 2019;Wang et al., 2020;Li et al., 2021;. The nuclear genome harbors biparentally inherited information and is essential to explore evolutionary events, such as hybridization and introgression (Rieseberg & Soltis, 1991;Soltis & Kuzoff, 1995;Ackerfield & Wen, 2003;Hardig et al., 2000;Bock et al., 2014). ...
Genome-scale data have significantly increased the number of informative characters for phylogenetic analyses and recent studies have also revealed widespread phylogenomic discordance in many plant lineages. Aralia sect. Aralia is a small plant lineage (14 spp.) of the ginseng family Araliaceae with a disjunct distribution between eastern Asia (11 spp.) and North America (3 spp.). We herein employ sequences of hundreds of nuclear loci and the complete plastomes using targeted sequence capture and genome skimming to reconstruct the phylogenetic and biogeographic history of this section. We detected substantial conflicts among nuclear genes, yet different analytical strategies generated largely congruent topologies from the nuclear data. Significant cytonuclear discordance was detected, especially concerning the positions of the three North American species. The phylogenomic results support two intercontinental disjunctions: (1) Aralia californica of western North America is sister to the eastern Asian clade consisting of A. cordata and A. continentalis in the nuclear tree, and (2) the eastern North American A. racemosa forms a clade with A. bicrenata from southwestern North America, and the North American A. racemosa - A. bicrenata clade is then sister to the eastern Asian clade consisting of A. glabra (Japan), A. fargesii (C China), and A. apioides and A. atropurpurea (the Hengduan Mountains). Aralia cordata is supported to be disjunctly distributed in Japan, Taiwan, the Ulleung island of Korea, and in Central, Southwest and South China, and Aralia continentalis is redefined with a narrower distribution in Northeast China, eastern Russia and peninsular Korea.
... Asteraceae is a cosmopolitan family divided into 13 subfamilies, 44 tribes (Panero et al. 2014), 1600 genera and 24000 species (Funk et al. 2009;Mandel et al. 2019). Anthemideae, which is one of the important tribes, consists of 111 genera and 1800 species worldwide. ...
A comparative anatomical study of the stems in 32 taxa assigned in the two related genera; Matricaria (4 taxa) and Tripleurospermum (28 taxa, 30 accessions), was carried out in Turkey by using cluster analysis (CA) and principal component analysis (PCA) to address generic classification and taxa delimitation. All the studied taxa have the following stem characteristics: one layer of epidermal cells with a sparse distribution of non-glandular trichomes, a cortex composed of interchanging collenchyma and chlorenchyma cells, a number of schizogenous secretory ducts near the bundles, ovate to oblong and open collateral vascular bundles which are arranged in a ring and are variable in size, and parenchymatous pith. The differences among the taxa are mainly the thickness of the xylem in the vascular bundle, the length of the epidermal cells and the pith cell size. The taxonomic and ecological values of the anatomical data are discussed in light of the current framework.
... Moreover, the radiate capitulum, which is a characteristic of most species in Asteroideae also occurred in the same subfamily. In this study, phylogenetic analysis was supported by high bootstrap values (BS > 98%) for morphological characteristics and phylogenetic relationships in Asteraceae (Panero and Funk 2002;Panero et al. 2014;Fu et al. 2016;Mandel et al. 2019;Zhang et al. 2021). Asteroideae is the largest subfamily of Astereaceae and contains 22 tribes (Fu et al. 2016). ...
Adenostemma madurense D.C. belongs to the family Asteraceae and is a wild annual herb found in Korea. In this study, we determined the phylogenetic origin of A. madurense by identifying its complete chloroplast sequence. The result indicated a genome size of 150,054 bp, which was composed of a large single copy of 82,008 bp, two inverted repeats of 24,952 bp each, and a small single-copy of 18,142 bp. Overall, 128 unique genes were identified, including 85 protein-coding, 35 tRNA, and 8 rRNA genes. A comparison of the A. madurense and A. lavenia chloroplast genomes revealed seven variations. Phylogenetic analysis revealed that A. madurense formed a clade with A. lavenia and had a well-supported phylogenetic relationship with Asteraceae. The first complete chloroplast genome of A. madurense was obtained, which would be helpful in explaining the process of speciation in Adenostemma and the development of molecular markers.
... On the basis of molecular studies, Asteraceae was divided into 10 subfamilies and 35 tribes (Baldwin et al. 2002;Panero and Funk 2002), although, for practical purposes, the most widely accepted classification is that of Bremer (1994), which includes 3 subfamilies and 17 tribes (Asteroideae, with 10 tribes; Barnadesioideae, with 1 tribe; and Cichorioideae, with 6 tribes). Currently, 44 tribes and 13 subfamilies of Asteraceae have been recognized based on phylogenetic studies (Panero et al. 2014). The number of palynological studies on the family is small, considering its taxonomic, economic, and numerical importance, but, since 2009, more attention has been given to the topic, as explained by Wortley et al. (2012). ...
Given the wide morphological variability of pollen grains and spores, palynology can provide important contributions to several branches of science. Palynological information, alone or in conjunction with other data, is particularly useful for the taxonomic delimitation of species, genera, families, and higher-rank taxa. The pollen character with the highest relevance as a taxonomic marker is the type of structure resulting from sporogenesis: if a spore, it characterizes the large groups of vascular (Pteridophyta s.l.) and avascular (Bryophyta s.l.) cryptogams; if a pollen grain, it characterizes gymno- and angiosperms. Pollen unit, polarity, aperture, and sexine ornamentation are other important pollen characters; these traits are genetically determined and do not respond to variations in environmental conditions. Palynology applied to taxonomy has been the major field of research of the Álvaro Xavier Moreira Laboratory of Palynology, Brazil. Palynotaxonomy has proven useful in the study of the families Asteraceae, Passifloraceae, Podostemaceae, Vitaceae, and Leguminosae, among others. It is noteworthy the growing use of palynology to support cladistic and multidisciplinary studies seeking to establish relationships and degrees of kinship between different groups of plants to trace the evolutionary history of taxa.
... A maximum-likelihood analysis based on the GTRGAMMA model was performed with Bayesian method on the CIPRES (Miller et al. 2010;Ronquist et al. 2012) using 1000 bootstrap replicates. The phylogenetic analysis of the cp genome dataset recovers the similar clades as in previous phylogenetic work (Panero and Funk 2008;Panero et al. 2014;Fu et al. 2016). The Bayesian inference (BI) result with 100% bootstrap showed that N. insignis has a close sister relationship with the genus Gerbrea (Figure 1). ...
This study was the first report complete chloroplast genome of Nouelia insignis (Asteraceae, Hyalideae), the large shrubs to small trees endemic to China. The circular whole cp genome of N. insignis was 151,524 bp in length, containing a large single-copy (LSC) region of 83,145 bp and a small single-copy (SSC) region of 18,261 bp. These two regions were separated by a pair of inverted repeat regions (IRa and IRb), each of them 25,060 bp in length. A total of 135 functional genes were encoded, consisting of 89 protein-coding genes, 38 tRNA genes, and eight rRNA genes. The overall GC content of the chloroplast genome sequence was 37.8%, and the GC contents of the LSC, SSC, and IR regions were 35.9, 31.5, and 43.2%, respectively. The phylogenetic analysis by the Bayesian analysis showed that the species of N. insignis was sister group with Gerbera jamesonii by strong support values, and thus was closely related to members of subfamilies of Cichorioideae and Pertyoideae. These results will be useful for the future studies of Asteraceae in the worldwide.
... The presence of pseudanthia is evolutionary labile with 95% confidence interval for total number of transitions reconstructed as 115-117 and at least 36 independent origins in a moderately large clade like Apioideae (Figure 2). Although limited, data available for several lineages of Asteraceae indicate that, in other plant groups, these structures are also prone to convergence (Francisco-Ortega et al., 1999;Sanz et al., 2008;Panero et al., 2014). The high evolvability of flower-like inflorescences can be tentatively explained by the developmental preadaptation and their inability to cause an irreversible phenotypical and ecological specialization, which would hinder eventual losses and re-gains. ...
Premise:
Pseudanthia are widespread and have long been postulated a key innovation responsible for some of the angiosperm radiations. The aim of our study was to analyze macroevolutionary patterns of these flower-like inflorescences and their potential correlation with diversification rates in Apiaceae subfamily Apioideae. In particular, we were interested to investigate evolvability of pseudanthia and evaluate their potential association with changes in size of floral display.
Methods:
The framework for our analyses consisted of time-calibrated phylogeny of 1734 representatives of Apioideae and a morphological matrix of inflorescence traits encoded for 847 species. Macroevolutionary patterns in pseudanthia were inferred using Markov models of discrete character evolution and stochastic character mapping. Additionally, a principal component analysis was conducted to visualize correlations in inflorescence architecture. The interdependence between net diversification rates and the occurrence of pseudocorollas was analysed with trait-independent and trait-dependent approaches.
Results:
Pseudanthia evolved in ten major clades of Apioideae with at least 36 independent origins and 46 reversals. The morphospace analysis recovered differences in color and compactness between floral and hyperfloral pseudanthia. A correlation between pseudocorollas and size of inflorescence was also strongly supported. Contrary to our predictions, pseudanthia are not responsible for variation in diversification rates identified in this subfamily.
Conclusions:
Our results suggest that pseudocorollas evolve as an answer to the trade-off between enlargement of floral display and costs associated with production of additional flowers. The high evolvability and architectural differences in apioid pseudanthia may be explained on the basis of adaptive wandering and evolutionary developmental biology. This article is protected by copyright. All rights reserved.
... La famille des Asteraceae (Compositae) compte environ 1600 genres et plus de 23000 espèces dans le monde, ce qui en fait l'une des plus vastes familles du règne végétal (Panero et al., 2014). Au Maghreb, cette famille comprend 512 genres et 1898 taxons, dont le Maroc et l'Algérie hébergent la moitié environ de la richesse générique (Dobignard & Chatelain, 2011). ...
The authors propose a state of knowledge about ecology and conservation issues of the
Algerian endemic Crepis arenaria (Pomel) Pomel subsp. arenaria, rediscovered after more
than half a century in the region of Aïn Sefra.
... Mais tarde Funk et al. (2009) aceitaram 1.700 gêneros subordinados a 12 subfamílias e 43 tribos. Panero et al. (2014), em um estudo filogenético para a família, criaram uma nova tribo e uma nova subfamília monotípica. ...
Os grãos de pólen de Asteraceae já vem sendo estudados há alguns anos no Laboratório de Palinologia Alvaro Xavier Moreira do Departamento de Botânica no Museu Nacional / UFRJ. É uma família rica de representantes com distribuição geográfica cosmopolita, ocorrendo em todos os continentes, com exceção do Antártico. No Parque Nacional de Itatiaia ocorrem cerca de 180 táxons de Asteraceae. Desse total foram analisados 109 espécies (ca. 60,0 %) distribuídas em 61 gêneros. Os táxons foram organizados em tribos, segundo a classificação mais aceita. Os grãos de pólen foram acetolizados segundo metodologia tradicional, medidos, fotomicrografados em microsocpia de luz. Para a obtenção das imagens em microscopia
eletrônica de varredura, os grãos de pólen não foram acetolizados. São ilustradas tanto as plantas quanto os grãos de pólen da maioria das espécies estudadas. São apresentadas descrições minuciosas dos principais atributos polínicos (forma, tamanho, abertura e ornamentação da sexina), tabelas onde estão registradas as medidas dos grãos de pólen em vista equatorial (diâmetros polar e equatorial). Entende-se que floras palinológicas e taxonômicas, são de grande valor para o conhecimento das espécies de uma determinada área, da distribuição e da morfologia dos táxons. Tendo em vista a grande diversidade das
Asteraceae e o alto valor da morfologia polínica de seus táxons, buscou-se caracterizar a morfologia polínica dos gêneros da família da área em estudo.
... Asteraceae is a cosmopolitan family, with a predominant occurrence in savanna and highland habitats (Hind 1993). Eupatorieae Cassini (1819: 202) is one of the 44 tribes circumscribed in the family (Panero et al. 2014) and includes about 180 genera and 2400 species (Robinson et al. 2009), being therefore considered one of the most diverse tribes within the family. Eupatorieae is a Neotropical tribe traditionally characterized by its opposite leaves, homogamous and discoid capitula, white to lilac tubular florets, long, clavate and sterile stylar branch appendages, and cypselae blackened due to phytomelanin deposition in the mature fruit (Robinson et al. 2009). ...
The southeastern Brazilian phytophysiognomy of “campos de altitude” is threatened by the global temperature increase, and because of it, many of its endemic species, such as those of the Asteraceae family, may disappear over the next years. This new species, which occurs in this type of environment, belongs to the genus Heterocondylus (Eupatorieae: Asteraceae), which comprises seven species, mainly with South American distribution. This genus is morphologically variable but all its species share the diagnostic characters, such as: enlarged basal style with variable indumentum, and cypselae with an asymmetrical carpopodium composed of cells with thickened walls. Heterocondylus penninervius is described as a new species from the “campos de altitude” in Serra do Brigadeiro, Araponga municipality, Minas Gerais state, Brazil. The species is recognized by its sessile leaves concentrated in the lower part of the stem, with narrow-elliptical to narrow-lanceolate blade and penninerved leaves, as well as broad capitula with white and pinkish phyllaries. This study provides the description, illustration, taxonomic affinities, flowering and fruiting period and geographic distribution of the species.
... The age of the root of the tree was calibrated with a uniform distribution of 73-101 Myr (maximum age of Asterales according to Beaulieu et al. 2013). The Asteraceae phylogeny does not completely agree with the phylogenetic relationships shown by Panero et al. (2014) despite using a starting tree, but the age estimates are congruent with other age estimates (Diplostephium and Linochilus clade by Vargas et al. (2018); Senecioneae by Kandziora et al. 2017; Euryops by Devos et al. 2010). We then used the resulting crown age estimate of the Erigeron-Diplostephium clade (7.7-22.06 ...
Páramo, the most species-rich tropical mountain ecosystem, is relatively well-researched in terms of the diversity and evolutionary sources of its flora, yet we know very little about the diversification within this environment. This study aims to unravel the evolutionary history of Oritrophium, an endemic genus of alpine habitats in North and South America, with a disjunct and bi-modal distribution of its species diversity. We aim to disentangle the center of origin and radiation of the genus, and mechanisms structuring its genetic diversity at inter-and intra-specific level. We sampled 19 species (85% from the total) and extended the sampling at population level for the two widely distributed species, O. limnophilum and O. peruvianum, comprising 19 and 24 populations, respectively. Using nuclear ribosomal internal transcribed spacer (ITS) and trnL-trnF chloroplast DNA region, we reconstructed dated phylogenies to test the monophyly of the genus and unravel possible historical forces underlying its diversification. We also performed an ancestral area estimation to reconstruct the biogeographic history of the genus. At the population level, we constructed haplotype networks and run spatial analyses of molecular variance to explore possible mechanisms that operate on structuring the diversity at intraspecific level. Oritrophium resulted polyphyletic, with two species being closely related to Erigeron and three other species ambiguously related to Erigeron, Diplostephium, Linochilus, and/or Hinterhubera. The remaining 14 species formed a clade, Oritrophium s.s., that likely originated during the Early Pliocene in the Andes of northwestern Bolivia to southern Ecuador, the center of the genus' diversity. The group likely diversified with the emergence of the Páramo during the Late Pliocene and further dispersed mainly from South-to-North in the Pleistocene. This migration involved both, long-distance dispersal from the Central Andes to Mexico and gradual migration of the species along the Andes. Accordingly, Oritrophium s.s. appears as the first record of a long-distance dispersal from the Páramo of South America to North America. The dispersal pattern within South America was mirrored by the intraspecific population diversity and structure of the investigated species.
... The Asteraceae-wide phylogeny was dated using an uncorrelated lognormal clock as implemented in BEAST v.2.6.2 (Bouckaert et al. 2014), based on two fossil calibration points (Barreda et al. 2012(Barreda et al. , 2015 and restricting the age of the root to 73-101 million years ago (Mya) (maximum age of the Asterales according to Beaulieu et al. 2013). The Asteraceae phylogeny does not completely agree with the phylogenetic relationships shown by Panero et al. (2014), despite using a starting tree, but the age estimates are congruent with other age estimates (Senecioneae, Kandziora et al. 2017;Euryops, Devos et al. 2010). The 95% confidence interval of the highest posterior density (HPD) values of the crown age of Dendrosenecio (16.77-2.07 ...
Alpine plant radiations are common across all major mountain systems of the world, and have been regarded as the main explanation for the species diversity found within these areas. To study the mechanisms behind the origin of this diversity, it is necessary to determine phylogenetic relationships and species boundaries in radiating alpine groups. The genus Dendrosenecio (Asteraceae) is an iconic example of a tropical-alpine plant radiation in the East African high mountains. To this date, limited sampling of molecular markers has resulted in insufficient phylogenetic resolution and infrageneric classification, hindering a comprehensive understanding of the drivers of diversification. Here, we used Hyb-Seq and the Compositae1061 probe set to generate targeted nuclear and off-target plastid DNA data for 42 samples representing all currently accepted 11 species. We combined coalescent methods and paralogy analysis to infer phylogenetic relationships, estimate divergence times and evaluate species boundaries. Lineage differentiation in Dendrosenecio seems to have occurred between the Late Miocene and the Pleistocene, starting when the first high elevation habitats became available in East Africa. We retrieved four major clades corresponding to four geographically distant mountain groups, testifying the importance of allopatric speciation in the early diversification of the group. Cytonuclear discordance suggested the occurrence of historical hybridization following occasional long-distance dispersal between mountain groups. The species delimitation analysis favored 10 species, but only five were fully supported, suggesting that population-level studies addressing processes such as ecological speciation and hybridization after secondary contact are needed to determine the current diversity found in the genus.
... These species are diverse in distributions and habitat, exist on every continent, including Antarctica, and occupied every type of habitat [1,2]. This family is divided into 13 subfamilies [1,3,4]. ...
The chloroplast genome evolves through the course of evolution. Various types of mutational events are found within the chloroplast genome, including insertions-deletions (InDels), substitutions, inversions, gene rearrangement, and pseudogenization of genes. The pseudogenization of the trnT-GGU gene was previously reported in the Cryptomeria japonica (Cupressaceae), Pelargonium x hortorum (Geraniaceae), and in the two species of the tribe Gnaphalieae (Asteroideae, Asteraceae). Here, we performed a broad analysis of the trnT-GGU gene among the species of twelve subfamilies of Asteraceae and found pseudogenization of this gene is not limited to the two species of Gnaphalieae or the tribe Gnaphalieae. We report for the first time that this gene is pseudo in the species of three subfamilies of Asteraceae, including Gymnarrhenoideae, Cichorioideae and Asteroideae. The analyses of the species of 78 genera of Asteroideae revealed that this pseudogenization event is linked to the insertion within the 5′ acceptor stem and not linked to the habit, habitat, and geographical distribution of the plant.
... Asteraceae (Compositae) is one of the largest family of flowering plants, comprises approximately 1,623 genera and 24,700 species (Christenhusz & Byng, 2016), belonging to 13 subfamilies and 44 tribes (Funk et al., 2009;Panero et al., 2014). The family has a cosmopolitan distribution throughout the world in temperate and tropical regions, occupying almost all the habitats (Funk et al., 2005). ...
THIS STUDY aimed to characterize the external morphology of achenes of the 74 taxa belonging to 51 genera and four subfamilies of Asteraceae from the Northwestern Mediterranean coast of Egypt. By using light microscopy, 26 quantitative and qualitative morphological characters were recorded. Among the studied taxa, achene symmetry, hairiness, surface topography, hilum position, presence or absence of pappus and pappus type were found significant diagnostic characters for delimiting the taxa. Two main groups were identified based on the presence or absence of hardened fruiting involucre. Four types and four subtypes were recognized: (1) Achene di- or trimorphic per capitulum, (2) Achene monomorphic and lacking pappus, (3) Achene monomorphic, with coroniform or auriculate pappus, (4) Achene monomorphic, with pappus. A description, photographs and identification keys were provided to assist in identification. Despite the taxonomic significance of achene exomorphology characters on the generic and specific level for the recognition of the different studied taxa of Asteraceae, no mutual link was shown between the taxonomic division of the family into subfamilies and tribes except for few cases.
... Targeted regions include the bi-parentally inherited whole nuclear ribosomal cistron, including the internal, external, and nontranscribed spacers, and the non-recombinant, uniparentally inherited chloroplast genome. Both nuclear ribosomal DNA (nrDNA) and chloroplast DNA (cpDNA) have been important in the reconstruction of the evolutionary history of Compositae at various scales of taxonomic organization (e.g., Baldwin et al., 1991;Susanna et al., 1995;Baldwin et al., 2002;Panero & Funk, 2008;Panero et al., 2014), but they often pose challenges to systematists when conflicts arise between trees generated from different sources of data (Baldwin, 1997;Vargas et al., 2017;Pouchon et al., 2018). Cytonuclear incongruence is common in Compositae and can be caused by error in phylogenetic inference, ongoing gene flow among lineages, hybridization, and incomplete lineage sorting (ILS) (Rieseberg & Soltis, 1991;Maddison, 1997;Huelsenbeck et al., 2000;Vargas et al., 2017). ...
Rock daisies (Perityleae; Compositae) are a diverse clade of seven genera and ca. 84 minimum‐rank taxa that mostly occur as narrow endemics on sheer rock‐cliffs throughout the southwest U.S. and northern Mexico. Taxonomy of Perityleae has traditionally been based on morphology and cytogenetics. To test taxonomic hypotheses and utility of characters emphasized in past treatments, we present the first densely sampled molecular phylogenies of Perityleae and reconstruct trait and chromosome evolution. We inferred phylogenetic trees from whole chloroplast genomes, nuclear ribosomal cistrons, and hundreds of low‐copy nuclear genes using genome skimming and target‐capture. Discordance between sources of molecular data suggests a underappreciated history of hybridization in Perityleae. Phylogenies support the monophyly of subtribe Peritylinae, a distinctive group possessing a four‐lobed disc corolla; however, all of the phylogenetic trees generated in this study reject the monophyly of the most species‐rich genus, Perityle, as well as its sections: Perityle sect. Perityle, Perityle sect. Laphamia, and Perityle sect. Pappothrix. Using reversible jump MCMC, our results suggest that morphological characters traditionally used to classify members of Perityleae have evolved multiple times within the group. A base chromosome number of x=9 gave rise to higher base numbers in subtribe Peritylinae (x=12, 13, 16, 17, 18 and 19) through polyploidization followed by ascending or descending dysploidy. Most taxa constitute a monophyletic lineage with a base chromosome number of x=17, with multiple neo‐polyploidization events. These results demonstrate the advantages and obstacles to next‐generation sequencing approaches in synantherology while laying the foundation for taxonomic revision and comparative study of the evolutionary ecology of Perityleae.
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... Asteraceae compreende 13 subfamílias, 44 tribos, cerca de 1.700 gêneros e aproximadamente 27.000 espécies, representando cerca de 10% do total das Angiospermas (Funk et al. 2009;Panero et al. 2014). No Brasil, há aproximadamente 2.097 espécies agrupadas em 290 gêneros (Flora do Brasil 2020, em construção). ...
Verbesina is an American genus of about 300 species with greater diversity in Mexico and the Andes. It is constituted by subshrubs and tree representatives, of alternate to opposite leaves, with entire to pinatipartids blades, capitulescence corymboid, capitulae discoid or radiated and cypselas winged and aristate.The objectives of this study were to perform the survey of the Brazilian Verbesina species, evaluate their circumscriptions and contribute to their knowledge through the presentation of descriptions, illustrations, identification key and ecological and taxonomic comments. The field survey and the study of herbarium specimens allowed the recognition of nine species for Brazil, occurring throughout the country, except for the North region, in environments of riparian and altitude forests. Five of the nine species of Verbesina (V. baccharifolia, V. bipinnatifida, V. floribunda, V. luetzelburgii and V. nicotianifolia) are exclusive to Brazil, with V. baccharifolia and V. luetzelburgii endemic to Bahia. Verbesina bipinnatifida is recorded for the first time for the Espírito Santo and V. subdiscoidea for Mato Grosso do Sul.
... The precise evolutionary relationships among the Fab Five tribes remain uncertain, with Senecioneae historically difficult to place & Funk, 2008;Panero et al., 2014;McDonald-Spicer et al., 2019), or as an unresolved polytomy ( Fig. 1D; Fu et al., 2016). In contrast, relationships among the four Asterodae tribes (Fu et al., 2016;Panero & Crozier, 2016) appear fairly well resolved and stable with Calenduleae (when sampled) as the first diverging lineage sister to a clade of Gnaphalieae, which, in turn, is sister to Astereae and Anthemideae (Fig. 1E). ...
Asteraceae account for 10% of all flowering plant species, and 35‐40% of these are in five closely‐related tribes that total over 10,000 species. These tribes include Anthemideae, Astereae, Calenduleae, Gnaphalieae, and Senecioneae, which form one of two enormous clades within Subfamily Asteroideae. We took a phylogenomics approach to resolve evolutionary relationships among these five tribes. We sampled the nuclear and plastid genomes via HybSeq target enrichment and genome skimming, and recovered 74 plastid genes and nearly 1000 nuclear loci, known as Conserved Orthologous Sequences. We tested for conflicting support in both datasets and used network analyses to assess patterns of reticulation to explain the early evolutionary history of this lineage which has experienced whole genome duplications and rapid radiations. We found both concordance and conflicting support in both datasets and documented four ancient hybridization events. Due to the timing of the early radiation of this five‐tribe lineage, shortly before the Eocene‐Oligocene extinction event (34 MYA), early lineages were likely lost, obscuring some details of their early evolutionary history.
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... For our MAPS analysis, we used transcriptomes from five Compositae species and four outgroup species each representing closely related families of the Compositae. Our species tree for the MAPS analysis was based on previously published phylogenies (Kim et al., 2005;Panero and Funk, 2008;Funk et al., 2009;Tank and Donoghue, 2010;Soltis et al., 2011;Panero et al., 2014) . ...
Premise of the study
Like many other flowering plants, members of the Compositae (Asteraceae) have a polyploid ancestry. Previous analyses found evidence for an ancient duplication or possibly triplication in the early evolutionary history of the family. We sought to better place this paleopolyploidy in the phylogeny and assess its nature.
Methods
We sequenced new transcriptomes for Barnadesia , the lineage sister to all other Compositae, and four representatives of closely related families. Using a recently developed algorithm, MAPS, we analyzed nuclear gene family phylogenies for evidence of paleopolyploidy.
Key results
We found that the previously recognized Compositae paleopolyploidy is also in the ancestry of the Calyceraceae. Our phylogenomic analyses uncovered evidence for a successive second round of genome duplication among all sampled Compositae except Barnadesia .
Conclusions
Our analyses of new samples with new tools provide a revised view of paleopolyploidy in the Compositae. Together with results from a high density Lactuca linkage map, our results suggest that the Compositae and Calyceraceae have a common paleotetraploid ancestor and most Compositae are descendants of a paleohexaploid. Although paleohexaploids have been previously identified, this is the first example where the paleotetraploid and paleohexaploid lineages have survived over tens of millions of years. The complex polyploidy in the ancestry of the Compositae and Calyceraceae represents a unique opportunity to study the long-term evolutionary fates and consequences of different ploidal levels.
... Asteraceae is the largest flowering plant family, comprises 23,000 species distributed in 13 subfamilies, 44 tribes and over 1600 genera in the World (Funk et al. 2009, Panero et al. 2014. This family distributes naturally on all continents outside of Antarctica, and its phylogenetic origin is thought to be in South America (Heywood 1978, Bremer 1994. ...
In this study, root, stem, leaf midrib and leaf lamina anatomy and achene micromorphology of Turkish critically endangered endemic Achillea sivasica were investigated for the first time. In this study, the root was found in late primary growth and in early secondary growth stage. It has large cortex layer consisting of 12-16 cell rows beneath the periderm. Secretory ducts formed by 5-12 secretory cells embedded in the cortex and located near vascular bundle were found at the root that is in early stage of secondary development. The stem was circular-pentagonal shaped in cross-section. There was lamellar collenchyma beneath epidermis of pentagon corners, and cortex parenchyma between corners. Secretory ducts located near the phloem, between the cortex and endodermis on the interfascicular region, were also observed. Endodermis layer was evident and its cells have indentations and protrusions where they touch adjacent endodermis cells, which strengthen the connection between them. In addition, casparian strips was conspicuous in many endodermis cells. Leaf midrib area was triangular shaped in cross section. There were secretory ducts, consisting of 4-5 secretory cells observed on both sides of the sclerenchymatous fibers which accompaning xylem. The leaf lamina was amphistomatic and stomata type was anomocytic. Mesophyll layer was equifacial. There was a large secretory duct and its diameter is bigger than the nearest main lamina vascular bundle. Achene shape was lanceolate-oblong and its surface was ribbed and glabrous of A. sivasica.
... For ycf1, we evaluated the potential utility of the Asteraceae ycf1 locus by reconstructing a phylogenetic tree based on the coding sequences of 211 ycf1 genes from Asteraceae whole plastomes in the NCBI nucleotide database. A ML tree constructed using the ycf1 genes showed that the Asteraceae species were divided into nine groups; in agreement with the tribe level taxonomic classification of the Asteraceae (Additional file 1: Fig. S4) [47][48][49]. The topology of a ML tree based on 212 Asteraceae accD genes was similar (Additional file 1: Fig. S5). ...
Background:
Artemisia in East Asia includes a number of economically important taxa that are widely used for food, medicinal, and ornamental purposes. The identification of taxa, however, has been hampered by insufficient diagnostic morphological characteristics and frequent natural hybridization. Development of novel DNA markers or barcodes with sufficient resolution to resolve taxonomic issues of Artemisia in East Asia is significant challenge.
Results:
To establish a molecular basis for taxonomic identification and comparative phylogenomic analysis of Artemisia, we newly determined 19 chloroplast genome (plastome) sequences of 18 Artemisia taxa in East Asia, de novo-assembled and annotated the plastomes of two taxa using publicly available Illumina reads, and compared them with 11 Artemisia plastomes reported previously. The plastomes of Artemisia were 150,858-151,318 base pairs (bp) in length and harbored 87 protein-coding genes, 37 transfer RNAs, and 8 ribosomal RNA genes in conserved order and orientation. Evolutionary analyses of whole plastomes and 80 non-redundant protein-coding genes revealed that the noncoding trnH-psbA spacer was highly variable in size and nucleotide sequence both between and within taxa, whereas the coding sequences of accD and ycf1 were under weak positive selection and relaxed selective constraints, respectively. Phylogenetic analysis of the whole plastomes based on maximum likelihood and Bayesian inference analyses yielded five groups of Artemisia plastomes clustered in the monophyletic subgenus Dracunculus and paraphyletic subgenus Artemisia, suggesting that the whole plastomes can be used as molecular markers to infer the chloroplast haplotypes of Artemisia taxa. Additionally, analysis of accD and ycf1 hotspots enabled the development of novel markers potentially applicable across the family Asteraceae with high discriminatory power.
Conclusions:
The complete sequences of the Artemisia plastomes are sufficiently polymorphic to be used as super-barcodes for this genus. It will facilitate the development of new molecular markers and study of the phylogenomic relationships of Artemisia species in the family Asteraceae.
... tonkinensis. These two regions have also been shown to be very efficient at distinguishing between different, closely related species belonging to multiple taxonomic groups, including bacteria and species from several different plant families (Kaneko, 2000;Iwasaki et al., 2012;Panero et al., 2014). The mutations in these two This article is protected by copyright. ...
Dalbergia odorifera T. C. Chen (Leguminosae), a rare and endangered tree species endemic to Hainan Island of China, produces the most expensive and rarest wood in China. The wood characteristics of D. odorifera are remarkably similar to those of D. tonkinensis (a much less sought‐after species from Vietnam), and the DNA from wood are often highly degraded, making it very difficult to identify the two species using anatomical features or DNA barcoding based on regular DNA markers. To solve the confusion of identifying wood reliably from the two species, we built and analyzed the plastome library of 26 samples from 18 Dalbergia species, of which 12 samples from eight close‐related species of D. odorifera are newly sequenced in this study. Phylogenomic analysis suggested that the relationships among the 26 samples are mostly well resolved, and conspecific individuals from different populations of D. odorifera and D. tonkinensis clustered together respectively. Between the plastid genomes of the two species, we identified 129 indels and 114 SNPs. By assessing a subset of 20 SNPs and ten indels using 37 population‐level samples (20 samples of D. odorifera and 17 samples of D. tonkinensis), we recovered eight species‐specific barcode regions which could be suitable for identifying the wood D. odorifera and D. tonkinensis. To examine their utility in wood identification, we amplified the eight DNA barcodes using six wood samples and recovered an amplification success rate of 83.3%, demonstrating a reliable method for precise wood identification of the two species.
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Convergent morphological evolution is widespread in flowering plants, and understanding this phenomenon relies on well-resolved phylogenies. Nuclear phylogenetic reconstruction using transcriptome datasets has been successful in various angiosperm groups, but it is limited to taxa with available fresh materials. Asteraceae are one of the two largest angiosperm families and important for both ecosystems and human livelihood, having multiple examples of convergent evolution. Nuclear Asteraceae phylogenies have resolved relationships among most subfamilies and many tribes, but many phylogenetic and evolutionary questions regarding subtribes and genera remain due to limited sampling. Here we increased the sampling for Asteraceae phylogenetic reconstruction using transcriptomes and genome skimming datasets and produced nuclear phylogenetic trees with 706 species representing two thirds of the recognized subtribes. Ancestral character reconstruction supports multiple convergent evolutionary events in Asteraceae, with gains and losses of bilateral floral symmetry correlated to the diversification of some subfamilies and smaller groups, respectively. The presence of the calyx-related pappus might have been especially important for the success of some subtribes and genera. Molecular evolutionary analyses supporting likely contribution of duplications of MADS-box and TCP floral regulatory genes to floral morphological innovations, including the capitulum inflorescence and bilaterally symmetric flowers, potentially promoting the diversification of Asteraceae. Subsequent divergences and reductions of CYC2 gene expression are related to the gain and loss of zygomorphic flowers. The phylogenomic work with greater taxon sampling by including GS datasets reveals the feasibility of expanded evolutionary analyses using DNA samples in understanding convergent evolution.
Parthenium hysterophorus , a globally widespread weed, poses a significant threat to agricultural ecosystems due to its invasive nature. We investigated the chloroplast genome of P. hysterophorus in this study. Our analysis revealed that the chloroplast genome of P. hysterophorus spans a length of 151,881 base pairs (bp). It exhibits typical quadripartite structure commonly found in chloroplast genomes, including inverted repeat regions (IR) of 25,085 bp, a small single copy (SSC) region of 18,052 bp, and a large single copy (LSC) region of 83,588 bp. A total of 129 unique genes were identified in P. hysterophorus chloroplast genomes, including 85 protein-coding genes, 36 tRNAs, and eight rRNAs genes. Comparative analysis of the P. hysterophorus plastome with those of related species from the tribe Heliantheae revealed both conserved structures and intriguing variations. While many structural elements were shared among the species, we identified a rearrangement in the large single-copy region of P. hysterophorus . Moreover, our study highlighted notable gene divergence in several specific genes, namely mat K, ndh F, clp P, rps 16, ndh A, rps 3, and ndh D. Phylogenetic analysis based on the 72 shared genes placed P. hysterophorus in a distinct clade alongside another species, P. argentatum . Additionally, the estimated divergence time between the Parthenium genus and Helianthus (sunflowers) was approximately 15.1 million years ago (Mya). These findings provide valuable insights into the evolutionary history and genetic relationships of P. hysterophorus , shedding light on its divergence and adaptation over time.
Angiosperms (flowering plants) are by far the most diverse land plant group with over 300,000 species. The sudden appearance of diverse angiosperms in the fossil record was referred to by Darwin as the “abominable mystery,” hence contributing to the heightened interest in angiosperm evolution. Angiosperms display wide ranges of morphological, physiological, and ecological characters, some of which have probably influenced their species richness. The evolutionary analyses of these characteristics help to address questions of angiosperm diversification and require well resolved phylogeny. Following the great successes of phylogenetic analyses using plastid sequences, dozens to thousands of nuclear genes from next‐generation sequencing have been used in angiosperm phylogenomic analyses, providing well resolved phylogenies and new insights into the evolution of angiosperms. In this review we focus on recent nuclear phylogenomic analyses of large angiosperm clades, orders, families, and subdivisions of some families and provide a summarized Nuclear Phylogenetic Tree of Angiosperm Families. The newly established nuclear phylogenetic relationships are highlighted and compared with previous phylogenetic results. The sequenced genomes of Amborella, Nymphaea, Chloranthus, Ceratophyllum, and species of monocots, Magnoliids, and basal eudicots, have facilitated the phylogenomics of relationships among five major angiosperms clades. All but one of the 64 angiosperm orders were included in nuclear phylogenomics with well resolved relationships except the placements of several orders. Most families have been included with robust and highly supported placements, especially for relationships within several large and important orders and families. Additionally, we examine the divergence time estimation and biogeographic analyses of angiosperm on the basis of the nuclear phylogenomic frameworks and discuss the differences compared with previous analyses. Furthermore, we discuss the implications of nuclear phylogenomic analyses on ancestral reconstruction of morphological, physiological, and ecological characters of angiosperm groups, limitations of current nuclear phylogenomic studies, and the taxa that require future attention.
This report aims to provide a framework for the efficient and effective ex situ conservation of globally important collections of sunflower and its crop wild relatives.
Background:
Pseudanthia or 'false flowers' are multiflowered units that resemble solitary flowers in form and function. Over the last century the term 'pseudanthium' has been applied to a wide array of morphologically divergent blossoms, ranging from those with easily noticeable florets to derived, reduced units in which individual flowers become almost indistinguishable. And although initially admired mostly by botanists, the diversity and widespread distribution of pseudanthia across angiosperms has already made them a fascinating topic for evolutionary and developmental comparative studies.
Scope:
This review synthesizes historical and current concepts on the biology of pseudanthia. Our first aim is to establish a clear, operational definition of pseudanthium and disentangle common terminological misconceptions surrounding that term. Our second aim is to summarize the knowledge of morphological and developmental diversity of pseudanthia and embed it within a modern phylogenetic framework. Lastly, we want to provide a comprehensive overview on the evolution and ecological importance of pseudanthia and outline perspectives for future studies.
Conclusions:
The understanding of pseudanthia changed multiple times and reflects three different interpretations of their 'flower-like' qualities: developmental (similarity in structure), figural (similarity in form and function), and phylogenetic (homology between angiosperm flowers and monoecious reproductive shoots in gymnosperms). Here, we propose to narrow the term pseudanthium to multiflowered blossoms resembling zoophilous flowers in form, i.e., in being structurally subdivided in a showy periphery and a reproductive center. According to this definition, pseudanthia sensu stricto. evolved independently in at least 40 angiosperm families. The recurrent acquisition of pseudanthia sensu stricto in all major lineages of flowering plants indicates repeated interactions between developmental constraints (smallness of flowers, meristematic conditions) and selective pressures, such as demands of pollinators and/or environmental conditions.
With c. 24 700 species (10% of all flowering plants), Asteraceae are one of the largest and most phenotypically diverse angiosperm families, with considerable economic and ecological importance. Asteraceae are distributed worldwide, from nearly polar latitudes all the way to the tropics, and occur across a diverse range of habitats from extreme deserts to swamps and from lowland rainforests to alpine tundra. Altogether, these characteristics make this family an outstanding model system to address a broad range of eco-evolutionary questions. In this review, we summarize recent progress in our understanding of Asteraceae on the basis of joint efforts by specialists in the fields of palaeobotany, cytogenetics, comparative genomics and phylogenomics. We will highlight how these developments are opening up new possibilities for integrating fields and better comprehending evolution beyond Asteraceae.
Artemisia lanaticapitula (Asteraceae: Anthemideae), a new species from Zhejiang in East China, is described and illustrated, based on the analyses of morphological observations and molecular phylogenetic evidence. This new species is similar to Artemisia anomala in having simple and undivided leaves, but differs by having phyllaries, corolla limbs, and abaxial leaf surfaces densely white-lanate, an involucre that is semiglobose, 3.5‐6 mm in diameter, and capitulate pedunculate at the base. It grows along streams under forests or on roadsides at forest margins; the species is narrowly distributed in Eastern and Southern Zhejiang.
The coastal landforms of Rio Grande do Sul originated very recently, and cover a wide range of distinct physiognomies which house a unique biodiversity, which is increasingly threatened. In this way, the Itapeva State Park is fundamental in the protection of the physiognomies and species characteristic of the coast. Considering that Asteraceae is the richest family in species number occurring in Rio Grande do Sul and one of the defining components of grasslands, this study aimed to produce a floristic list of the species of Asteraceae found in Itapeva State Park. This is the first study focused on listing solely species of Asteraceae conducted in the Park. Through field expeditions, use of the management plan and revisions of herbaria collections, we listed a total of 154 species, which include threatened species of genera Baccharis, Mikania, Noticastrum and Porophyllum, as well as 36 new occurrences to the Park. Differences between the species lists obtained through the different methods are attributed to the dynamics of the coastal landforms and the impact caused by tourism. Our results reaffirm the importance of floristic studies and of Asteraceae to the flora of Rio Grande do Sul. RESUMO As formações litorâneas do Rio Grande do Sul, de origem muito recente, abrangem uma ampla gama de fisionomias distintas e abrigam uma biodiversidade única e cada vez mais ameaçada. Neste sentido, o Parque Estadual de Itapeva é instrumental na proteção das fisionomias e espécies típicas do litoral. Sendo Asteraceae a família mais rica em espécies no estado e um importante componente das fisionomias campestres, este estudo objetivou produzir uma lista florística das espécies da família ocorrentes no PE de Itapeva. Este é o primeiro estudo realizado no local focado em listar apenas espécies de Asteraceae. Através de expedições a campo, consulta ao plano de manejo e a coleções de herbários, foi registrado um total de 154 espécies, incluindo espécies ameaçadas de Baccharis, Mikania, Noticastrum e Porophyllum e 36 novas ocorrências para o parque. Discrepâncias entre as listas obtidas através dos diferentes meios são atribuídas principalmente à dinâmica dos ambientes costeiros e ao impacto causado pelo turismo. Os resultados obtidos reiteram a importância da realização de estudos florísticos e da família Asteraceae para a flora do estado.
Palavras-chave: Compositae, levantamento de espécies, região litorânea
The necrotrophic fungal pathogen Sclerotinia sclerotiorum can cause disease on numerous plant species, including many important crops. Most S. sclerotiorum-incited diseases of crop plants are initiated by airborne ascospores produced when fungal sclerotia germinate to form spore-bearing apothecia. However, basal stalk rot of sunflower occurs when S. sclerotiorum sclerotia germinate to form mycelia within the soil which subsequently invade sunflower roots. To determine if other plant species in the Asteraceae family are susceptible to root infection by S. sclerotiorum, cultivated sunflower (Helianthus annuus L.) and seven other Asteraceae species were evaluated for S. sclerotiorum root infection by inoculation with either sclerotia or mycelial inoculum. Additionally, root susceptibility of sunflower was compared to that of dry edible bean and canola, two plant species susceptible to S. sclerotiorum but not known to display root-initiated infections. Results indicated that multiple Asteraceae family plants are susceptible to S. sclerotiorum root infection after inoculation with either sclerotia or mycelium. These observations expand the range of plant hosts susceptible to S. sclerotiorum root infection, elucidate differences in root inoculation methodology, and emphasize the importance of soil-borne infection to Asteraceae crop and weed species.
The chloroplast genome evolves through the course of evolution. Various types of mutational events are found within the chloroplast genome, including insertions-deletions (InDels), substitutions, inversions, gene rearrangement, and pseudogenization of genes. The pseudogenization of the chloroplast threonine (trnT-GGU) gene was previously reported in Cryptomeria japonica (Cupressaceae), Pelargonium × hortorum (Geraniaceae), and Anaphalis sinica and Leontopodium leiolepis of the tribe Gnaphalieae (Asteroideae, Asteraceae). Here, we performed a broad analysis of the trnT-GGU gene among the species of 13 subfamilies of Asteraceae and found this gene as a pseudogene in core Asteraceae (Gymnarrhenoideae, Cichorioideae, Corymbioideae, and Asteroideae), which was linked to an insertion event within the 5′ acceptor stem and is not associated with ecological factors such as habit, habitat, and geographical distribution of the species. The pseudogenization of trnT-GGU was not predicted in codon usage, indicating that the superwobbling phenomenon occurs in core Asteraceae in which a single transfer RNA (trnT-UGU) decodes all four codons of threonine. To the best of our knowledge, this is the first evidence of a complete clade of a plant species using the superwobbling phenomenon for translation.
We assembled new plastomes of 19 species of Mikania and of Ageratina fastigiata, Litothamnus nitidus, and Stevia collina, all belonging to tribe Eupatorieae (Asteraceae). We analyzed the structure and content of the assembled plastomes and used the newly generated sequences to infer phylogenetic relationships and study the effects of different data partitions and inference methods on the topologies. Most phylogenetic studies with plastomes ignore that processes like recombination and biparental inheritance can occur in this organelle, using the whole genome as a single locus. Our study sought to compare this approach with multispecies coalescent methods that assume that different parts of the genome evolve at different rates. We found that the overall gene content, structure, and orientation are very conserved in all plastomes of the studied species. As observed in other Asteraceae, the 22 plastomes assembled here contain two nested inversions in the LSC region. The plastomes show similar length and the same gene content. The two most variable regions within Mikania are rpl32-ndhF and rpl16-rps3, while the three genes with the highest percentage of variable sites are ycf1, rpoA, and psbT. We generated six phylogenetic trees using concatenated maximum likelihood and multispecies coalescent methods and three data partitions: coding and non-coding sequences and both combined. All trees strongly support that the sampled Mikania species form a monophyletic group, which is further subdivided into three clades. The internal relationships within each clade are sensitive to the data partitioning and inference methods employed. The trees resulting from concatenated analysis are more similar among each other than to the correspondent tree generated with the same data partition but a different method. The multispecies coalescent analysis indicate a high level of incongruence between species and gene trees. The lack of resolution and congruence among trees can be explained by the sparse sampling (~ 0.45% of the currently accepted species) and by the low number of informative characters present in the sequences. Our study sheds light into the impact of data partitioning and methods over phylogenetic resolution and brings relevant information for the study of Mikania diversity and evolution, as well as for the Asteraceae family as a whole.
In the present study light and scanning electron microscopic approaches were used to study the palynological variations among 30 Asteroideae members. Variety of qualitative and quantitative palynological characters like pollen shape and type, pollen sculpturing, pollen size, P/E ratio, exine and intine thickness, interspecific difference, colpi size, pore size, spines length, spine width, and pollen fertility were observed. Results reported remarkable variations among pollen characters of studied taxa. The shape of pollen grains in polar and equatorial views varied from specie to specie such as spheroidal, prolate, oblate, subspheroidal, oblate spheroidal, prolate spheroidal, suboblate, and sub prolate. Bidens pilosa L. showed maximum pollen size in both polar and equatorial views, whereas minimum pollen size in polar view was found in Tetraneuris scaposa (DC.) Greene. (i.e., 30.5 μm) and in equatorial view was found in Cosmos sulphureus Cav. (25.5 μm).The highest P/E ratio (1.39 μm) was measured in Glebionis coronaria L. The values of exine and intine thickness also varied. Tricolporate, tetracolporate, trizonocolporate, pentoporate, and polypentoporate pollen types were examined. Spinateand echinate pollen sculpturingobserved under SEM. Variations in spine length and width also recorded. The maximum pollen fertility (98%) was measured in Chrysanthemum morifolium Ramat. and the lowest (56%) was recorded in Thymophylla tenuiloba (DC.) Small. It was concluded that the variations in qualitative and quantitative characters were seemed to be valuable for the taxonomic investigations of Asteroideae taxa.
Osmophores are engaged in scent production and differ from other secretory structures by their product, site, duration, and anatomical structure. Whereas osmophores have been well-documented in flowers of several families, they are barely mentioned in the Asteraceae. The aims of this study are to: 1) determine the occurrence of osmophores in corollas of 39 species of the tribe Onoserideae and the only species of Famatinantheae with histochemical methods; and 2) analyze the morphology and structure of osmophores in these groups. Histochemical and histological techniques revealed osmophores in the marginal and in the central corollas of Famatinanthus Ariza & S.E. Freire and Plazia Ruiz & Pav. at the apex and margin of the corolla lobes, and at the sinuses. Osmophores were confirmed following positive staining reactions for TIOFH and TIOFH3 (Neutral Red, Oil Red O, and Iodine-Potassium-Iodide); the samples reacted negatively for Benedict's test for detecting reducing sugars. The osmophores found in this study are constituted uniquely by a papillose or non-papillose epidermis, or they extend into the mesophyll forming scent glands. In both cases, they are associated to stomata and starch grains. The corollas of the other species of Onoserideae did not react positively for osmophores.
Resumo Moquiniastrum, originalmente descrito por Cabrera como uma seção de Gochnatia, após análises moleculares e morfológicas mais detalhadas foi elevado à categoria de gênero. Moquiniastrum diferencia-se de Gochnatia pela presença de ginodioicia, indumento de tricomas com 2-5-raios e capitulescências paniculiformes. Atualmente é constituído por 21 espécies, distribuídas principalmente no Brasil. Pouco se conhece sobre a taxonomia do gênero Moquiniastrum e ainda há escassez de estudos sobre o arranjo sexual das flores no capítulo, necessitando de análises aprofundadas destas estruturas. O objetivo geral deste trabalho foi realizar um estudo taxonômico de Moquiniastrum na Região Sul do Brasil, além de avaliar e enquadrar os táxons em categorias de ameaça de extinção. Este estudo foi realizado através de revisões de herbários e expedições de campo nos três estados da Região Sul do Brasil. Foram encontradas 10 espécies de Moquiniastrum, além de três subespécies de M. polymorphum. São fornecidos chave de identificação para os táxons confirmados, descrições morfológicas, comentários sobre habitats, conservação e períodos de floração e de frutificação das espécies, além de observações taxonômicas e imagens.
The eudicots are a large, monophyletic assemblage of angiosperms, comprising roughly 190,000 described species, or 75% of all angiosperms. The monophyly of eudicots is well supported from molecular data and delimited by at least one palynological apomorphy: a tricolpate or tricolpate-derived pollen grain. A tricolpate pollen grain is one that has three apertures, equally spaced and approximately parallel to the polar axis of the grain. Apertures are differentiated regions of the pollen grain wall that may function as the site of pollen tube exitus as well as to allow for expansion and contraction of the pollen grain with changes in humidity. Tricolpate pollen grains evolved from a monosulcate type (having a single distal aperture, which is considered to be ancestral in the angiosperms, as well as for many seed plant clades. Many eudicots have pollen grains with more than three apertures, of a great variety of numbers, shapes, and position (constituting important taxonomic characters). These are all thought to have been derived from a tricolpate type.
In the present study anatomical characterization of 20 medicinally important Asteroideae species were done under light and scanning electron microscopy. Variety of qualitative and quantitative anatomical characters like epidermal cells, stomata, guard cells, subsidiary cells, trichomes and oil droplets were observed. Generally pentagonal, polygonal, irregular or hexagonal, smooth, undulating thick walled epidermal cells were observed in studied species. In abaxial surface Thymophylla tenuiloba L. possessed the largest length of epidermal cell that is, 221.6 (156.6–286.6) μm whereas Bellis perenis L. showed the smallest length that is, 46.4 (32.6–60.2) μm. Average width of epidermal cells ranged from 57 (22–92) μm to 169 (127.9–210.1) μm. Cosmos sulphureus Cav. had smallest width while Thymophylla tenuiloba L. had the largest width. In adaxial surface Artemisia absinthium L. possessed the largest length of epidermal cell that is, 269 (165.1–372.9) μm whereas Bellis perenis L. showed the smallest length that is, 61.4 (42.6–80.2) μm. Average width of epidermal cells ranged from 50.8 (32.6–69) μm to 260 (116–202) μm. Thymophylla tenuiloba L. had smallest width while Dahlia pinnata Cav. had the largest width. Among stomatal characters anisocytic, anomocytic, and diacytic stomata were observed in selected species of Asteroideae. Nonglandular uniserate, multicellular, unbranched pointed tips with bulbous base trichomes were reported in some Asteroideae members while some possessed glandular, capitates mushroom like multicellular trichomes covered with tubercle papicles. Rounded, oval, triangular shaped oil droplets were observed in some species. It is concluded that qualitative and qualitative anatomical variations in trichomes, stomata and epidermal cells are of good taxonomic value for the Asteroideae species.
The release of pollen in portions, i. e. limiting the pollen removed by individual pollinators during a single visit, is a widespread phenomenon. The biological significance of this phenomenon is that it helps to optimize the pollination. Parameters like number of ovules, number of application sites to the pollinator or spectrum of pollinators, which may be correlated with the size of the pollen portion, are illustrated as a net of possible correlations (Fig. 1). Apart from pollenkitt, viscin or cellulosic threads and the non-simultaneous opening of the anthers, the secondary pollen presentation is a special mechanism of pollen portioning. Secondary pollen presentation occurs in several dicotyledonous and in a few monocotyledonous families. Its occurrence is a distinctive feature in the Campanulales-Asterales-complex. The different mechanisms in the Campanulales-Asterales (Asteraceae, Lobeliaceae, Campanulaceae, Goodeniaceae, Brunoniaceae, Calyceraceae) are briefly described. The differentiations of the "basic syndrome" (radial symmetry of the flower bud, complete androecial whorl, introrse anthers, the latter connivent at the time of dehiscence, proterandry, late and sequential growth of filaments and style) are presented in two diagrams (Figs. 30 + 31).
At the Compositae Conference at Kew in 1994 ("Com-pos itae: Systematics, Biology, Utilization"), DeVore and Stuessy (1995) argued in favor of a sister group rela-tionship between Asteraceae and Calyceraceae, mainly drawing evidences from morphology. Even if a close relationship between Asteraceae and Calyceraceae had been repeatedly suggested ever since Cassini described the latter family (as Boopideae; Cassini 1816), it was perhaps not until the Compositae Conference at Reading in 1975 ("The Biology and Chemistry of the Com-positae"), that this relationship was seriously corrobo-rated (Turner 1977), mainly based on similarities in pol-len morphology (Skvarla et al. 1977). This hypothesis further gained strength with the identifi cation of the subtribe Barnadesiinae (now subfamily Barnadesioideae) in Mutisieae as the sister group to the rest of the fam-ily (Bremer 1987; Jansen and Palmer 1987; Bremer and Jansen 1992; Olmstead et al. 1992). However, the Asteraceae-Calyceraceae sister group relationship was soon challenged by Goodeniaceae (all these taxa, ex-cept Asteraceae of course, will be presented in some detail below). Since then there have been three com-peting hypotheses: a clade of Asteraceae + Calyceraceae with Goodeniaceae (if sampled) as its their sister group (Gustafsson and Bremer 1995; Kim and Jansen 1995; Downie et al. 1996; Jansen and Kim 1996; Bremer and Gustafsson 1997; Carlquist and DeVore 1998; Kårehed et al. 1999; Olmstead et al. 2000; Albach et al. 2001; K. Bremer et al. 2001; B. Bremer et al. 2002; Lundberg and Bremer 2003; Winkworth et al. 2008), or a clade of Goodeniaceae + Calyceraceae with Asteraceae as its sister group (Michaels et al. 1993; Olmstead et al. 1993; Cosner et al. 1994; Savolainen et al. 2000; Soltis et al. 2000, 2007), or a clade of Asteraceae + Goodeniaceae with Calyceraceae (if sampled) as its sister group (Gustafsson and Bremer 1995; Gustafsson et al. 1996). It is possible to fi nd at least some characters in favor of any of these relations (as well as contradicting them), but as shown by DeVore and Stuessy (1993), Hansen (1997), and Lundberg and Bremer (2003), the morphology is mainly in favor of the Calyceraceae-Asteraceae sister group relationship, while it is largely some molecular markers that suggested the other two alternatives. Furthermore, the two best-sampled analyses to date (Lundberg and Bremer 2003; Winkworth et al. 2008) both support the Calyceraceae-Asteraceae sister group relationship. This contribution does not argue for this sister group relationship, but instead gives an overview of what I think is the most likely phylogeny of the Asteraceae alliance, covering the entire order Asterales (sensu APG II 2003; Fig. 10.1; Table 10.1).
Abstract— Famatinanthus , a new genus of Asteraceae (Mutisioideae, Onoserideae), is described and illustrated to accommodate one species from the Andes of Argentina, that was previously placed in Aphyllocladus, A. decussatus, as Famatinanthus decussatus comb. nov. The new genus is tentatively assigned to the tribe Onoserideae based on its shrubby habit, solitary radiate capitula, style rounded at the apex and dorsally papillose, and 2‐3-seriate heteromorphic pappus. Famatinanthus is similar to Aphyllocladus but it is easily distinguished by the leafy, decussate branches with opposite leaves, multistoried T-trichomes, cream corollas, apiculate apical anther appendages, setuliferous achenes, terete stems, lack of secretory cavities, and pollen with a conspicuous mesoaperture and microechinate-rugulate exine. A key to the genera of the Onoserideae is presented. Affinities of the new genus with other genera of the tribes Gochnatieae, Hyalideae, and Stifftieae are also discussed.
• Premise of the study: The Compositae (Asteraceae) are a large and diverse family of plants, and the most comprehensive phylogeny to date is a meta-tree based on 10 chloroplast loci that has several major unresolved nodes. We describe the development of an approach that enables the rapid sequencing of large numbers of orthologous nuclear loci to facilitate efficient phylogenomic analyses.
• Methods and Results: We designed a set of sequence capture probes that target conserved orthologous sequences in the Compositae. We also developed a bioinformatic and phylogenetic workflow for processing and analyzing the resulting data. Application of our approach to 15 species from across the Compositae resulted in the production of phylogenetically informative sequence data from 763 loci and the successful reconstruction of known phylogenetic relationships across the family.
• Conclusions: These methods should be of great use to members of the broader Compositae community, and the general approach should also be of use to researchers studying other families.
The tempo of species diversification in large clades can reveal fundamental evolutionary mechanisms that operate on large temporal and spatial scales [1-4]. Hummingbirds have radiated into a diverse assemblage of specialized nectarivores comprising 338 species, but their evolutionary history has not, until now, been comprehensively explored. We studied hummingbird diversification by estimating a time-calibrated phylogeny for 284 hummingbird species, demonstrating that hummingbirds invaded South America by ∼22 million years ago, and subsequently diversified into nine principal clades (see [5-7]). Using ancestral state reconstruction and diversification analyses, we (1) estimate the age of the crown-group hummingbird assemblage, (2) investigate the timing and patterns of lineage accumulation for hummingbirds overall and regionally, and (3) evaluate the role of Andean uplift in hummingbird speciation. Detailed analyses reveal disparate clade-specific processes that allowed for ongoing species diversification. One factor was significant variation among clades in diversification rates. For example, the nine principal clades of hummingbirds exhibit ∼15-fold variation in net diversification rates, with evidence for accelerated speciation of a clade that includes the Bee, Emerald, and Mountain Gem groups of hummingbirds. A second factor was colonization of key geographic regions, which opened up new ecological niches. For example, some clades diversified in the context of the uplift of the Andes Mountains, whereas others were affected by the formation of the Panamanian land bridge. Finally, although species accumulation is slowing in all groups of hummingbirds, several major clades maintain rapid rates of diversification on par with classical examples of rapid adaptive radiation.
Background
New powerful biogeographic methods have focused attention on long-standing hypotheses regarding the influence of the break-up of Gondwana on the biogeography of Southern Hemisphere plant groups. Studies to date have often concluded that these groups are too young to have been influenced by these ancient continental movements. Here we examine a much larger and older angiosperm clade, the Campanulidae, and infer its biogeographic history by combining Bayesian divergence time information with a likelihood-based biogeographic model focused on the Gondwanan landmasses.
Results
Our analyses imply that campanulids likely originated in the middle Albian (~105 Ma), and that a substantial portion of the early evolutionary history of campanulids took place in the Southern Hemisphere, despite their greater species richness in the Northern Hemisphere today. We also discovered several disjunctions that show biogeographic and temporal correspondence with the break-up of Gondwana.
Conclusions
While it is possible to discern traces of the break-up of Gondwana in clades that are old enough, it will generally be difficult to be confident in continental movement as the prime cause of geographic disjunctions. This follows from the need for the geographic disjunction, the inferred biogeographic scenario, and the dating of the lineage splitting events to be consistent with the causal hypothesis.
The Goodeniaceae, close relatives of the Asteraceae, are a conspicuous part of the flora of Australia and many islands in the Pacific. A comprehensive molecular phylogenetic analysis of the family using cpDNA regions trnL-F and matK is presented, including representatives of all genera and nearly half the species. The family resolves into the two large clades: 'LAD' (Lechenaultia, Anthotium, Dampiera) and 'Core Goodeniaceae' (Brunonia, Scaevola, Diaspasis, Coopernookia, Goodenia, Selliera, Velleia, Verreauxia, Pentaptilon), which are also supported by morphological characters. The former Brunoniaceae, comprising the single species Brunonia australis, is clearly placed within the Goodeniaceae as sister to the remainder of Core Goodeniaceae while possessing many autapomorphic characteristics. Current subgeneric taxonomic groups are partially supported as monophyletic within Goodenia and Dampiera, but are generally non-monophyletic within Lechenaultia and Scaevola. Fruit types and floral traits involved in pollination and adaptations to aridity have evolved in a highly convergent fashion within several major clades. Rates of molecular evolution and number of extant taxa differ at several points between sister clades, particularly between Brunonia and the rest of Core Goodeniaceae, and between Scaevola s.l. and Goodenia s.l. Future taxonomic changes will involve the synonymization of Selliera and monotypic Diaspasis and, pending more comprehensive taxon and molecular sampling, the large, paraphyletic genus Goodenia may be split into at least three genera.
Understanding the evolutionary history of living organisms is a central problem in biology. Until recently the ability to infer evolutionary relationships was limited by the amount of DNA sequence data available, but new DNA sequencing technologies have largely removed this limitation. As a result, DNA sequence data are readily available or obtainable for a wide spectrum of organisms, thus creating an unprecedented opportunity to explore evolutionary relationships broadly and deeply across the Tree of Life. Unfortunately, the algorithms used to infer evolutionary relationships are NP-hard, so the dramatic increase in available DNA sequence data has created a commensurate increase in the need for access to powerful computational resources. Local laptop or desktop machines are no longer viable for analysis of the larger data sets available today, and progress in the field relies upon access to large, scalable high-performance computing resources. This paper describes development of the CIPRES Science Gateway, a web portal designed to provide researchers with transparent access to the fastest available community codes for inference of phylogenetic relationships, and implementation of these codes on scalable computational resources. Meeting the needs of the community has included developing infrastructure to provide access, working with the community to improve existing community codes, developing infrastructure to insure the portal is scalable to the entire systematics community, and adopting strategies that make the project sustainable by the community. The CIPRES Science Gateway has allowed more than 1800 unique users to run jobs that required 2.5 million Service Units since its release in December 2009. (A Service Unit is a CPU-hour at unit priority).
Morphological, molecular and biogeographical information bearing on early evolution of the sunflower alliance of families suggests that the clade containing the extant daisy family (Asteraceae) differentiated in South America during the Eocene, although palaeontological studies on this continent failed to reveal conclusive support for this hypothesis. Here we describe in detail Raiguenrayun cura gen. & sp. nov., an exceptionally well preserved capitulescence of Asteraceae recovered from Eocene deposits of northwestern Patagonia, Argentina.
The fossil was collected from the 47·5 million-year-old Huitrera Formation at the Estancia Don Hipólito locality, Río Negro Province, Argentina.
The arrangement of the capitula in a cymose capitulescence, the many-flowered capitula with multiseriate-imbricate involucral bracts and the pappus-like structures indicate a close morphological relationship with Asteraceae. Raiguenrayun cura and the associated pollen Mutisiapollis telleriae do not match exactly any living member of the family, and clearly represent extinct taxa. They share a mosaic of morphological features today recognized in taxa phylogenetically close to the root of Asteraceae, such as Stifftieae, Wunderlichioideae and Gochnatieae (Mutisioideae sensu lato) and Dicomeae and Oldenburgieae (Carduoideae), today endemic to or mainly distributed in South America and Africa, respectively.
This is the first fossil genus of Asteraceae based on an outstandingly preserved capitulescence that might represent the ancestor of Mutisioideae-Carduoideae. It might have evolved in southern South America some time during the early Palaeogene and subsequently entered Africa, before the biogeographical isolation of these continents became much more pronounced. The new fossil represents the first reliable point for calibration, favouring an earlier date to the split between Barnadesioideae and the rest of Asteraceae than previously thought, which can be traced back at least 47·5 million years. This is the oldest well dated member of Asteraceae and perhaps the earliest indirect evidence for bird pollination in the family.
Premise of the study:
Phylogenies based on molecular data are revealing that generalizations about complex morphological structures often obscure variation and developmental patterns important for understanding the evolution of forms, as is the case for inflorescence morphology within the well-supported MGCA clade (Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae). While the basal families share a basic thyrsic/thyrsoid structure of their inflorescences, Asteraceae possesses a capitulum that is widely interpreted as a racemose, condensed inflorescence. Elucidating the poorly known inflorescence structure of Calyceraceae, sister to Asteraceae, should help clarify how the Asteraceae capitulum evolved from thyrsic/thyrsoid inflorescences.
Methods:
The early development and structure of the inflorescence of eight species (five genera) of Calyceraceae were studied by SEM, and patterns of evolutionary change were interpreted via phylogenetic character mapping.
Key results:
The basic inflorescence structure of Calyceraceae is a cephalioid (a very condensed botryoid/thyrsoid). Optimization of inflorescence characters on a DNA sequence-derived tree suggests that the Asteraceae capitulum derives from a simple cephalioid through two morphological changes: loss of the terminal flower and suppression of the cymose branching pattern in the peripheral branches.
Conclusions:
Widely understood as a condensed raceme, the Asteraceae capitulum is the evolutionary result of a very reduced, condensed thyrsoid. Starting from that point, evolution worked separately only on the racemose developmental control/pattern within Asteraceae and mainly on the cymose developmental control/pattern within Calyceraceae, producing head-like inflorescences in both groups but with very different diversification potential. We also discuss possible remnants of the ancestral cephalioid structure in some Asteraceae.
Subfamily Barnadesioideae (Asteraceae) consists of nine genera and 91 species endemic to South America. They include annual and perennial herbs, arching shrubs and trees up to 30m tall. Presumed sister to all other Asteraceae, its intergeneric relationships are key to understanding the early evolution of the family. Results of the only molecular study on the subfamily conflict with relationships inferred from morphology. We investigate inter- and intrageneric relationships in Barnadesioideae with novel DNA sequence data and morphological characters using parsimony, likelihood and Bayesian inference. All results verify Barnadesioideae as monophyletic and sister to the rest of the family. A basal split within the subfamily is recognized, with Chuquiraga, Doniophyton and Duseniella in one clade, and Arnaldoa, Barnadesia, Dasyphyllum, Fulcaldea, Huarpea and possibly Schlechtendalia in another. The largest genus, Dasyphyllum, is revealed as biphyletic with the two clades separating along subgeneric and geographic lines. Schlechtendalia, suggested as the earliest diverging lineage of the subfamily by morphological studies and parsimony analyses, is found in a more derived position under model-based inference methods. Competing phylogenetic hypotheses, both previous and present, are evaluated using likelihood-based tests. Evolutionary trends within Barnadesioideae are inferred: hummingbird pollination has developed convergently at least three times. An early vicariance in the subfamily's distribution is revealed. X=9 is supported as the ancestral base chromosome number for both Barnadesioideae and the family as a whole.
An extensive sequence comparison of the chloroplast ndhF gene from all major clades of the largest flowering plant family (Asteraceae) shows that this gene provides approximately 3 times more phylogenetic information than rbcL. This is because it is substantially longer and evolves twice as fast. The 5' region (1380 bp) of ndhF is very different from the 3' region (855 bp) and is similar to rbcL in both the rate and the pattern of sequence change. The 3' region is more A+T-rich, has higher levels of nonsynonymous base substitution, and shows greater transversion bias at all codon positions. These differences probably reflect different functional constraints on the 5' and 3' regions of ndhF. The two patterns of base substitutions of ndhF are particularly advantageous for phylogenetic reconstruction because the conserved and variable segments can be used for older and recent groups, respectively. Phylogenetic analyses of 94 ndhF sequences provided much better resolution of relationships than previous molecular and morphological phylogenies of the Asteraceae. The ndhF tree identified five major clades: (i) the Calyceraceae is the sister family of Asteraceae; (ii) the Barnadesioideae is monophyletic and is the sister group to the rest of the family; (iii) the Cichorioideae and its two basal tribes Mutisieae and Cardueae are paraphyletic; (iv) four tribes of Cichorioideae (Lactuceae, Arctoteae, Liabeae, and Vernonieae) form a monophyletic group, and these are the sister clade of the Asteroideae; and (v) the Asteroideae is monophyletic and includes three major clades.
Evolutionary biologists have adopted simple likelihood models for purposes of estimating ancestral states and evaluating character
independence on specified phylogenies; however, for purposes of estimating phylogenies by using discrete morphological data,
maximum parsimony remains the only option. This paper explores the possibility of using standard, well-behaved Markov models
for estimating morphological phylogenies (including branch lengths) under the likelihood criterion. An important modification
of standard Markov models involves making the likelihood conditional on characters being variable, because constant characters
are absent in morphological data sets. Without this modification, branch lengths are often overestimated, resulting in potentially
serious biases in tree topology selection. Several new avenues of research are opened by an explicitly model-based approach
to phylogenetic analysis of discrete morphological data, including combined-data likelihood analyses (morphology + sequence
data), likelihood ratio tests, and Bayesian analyses.
MrBayes 3 performs Bayesian phylogenetic analysis combining information from different data partitions or subsets evolving under different stochastic evolutionary models. This allows the user to analyze heterogeneous data sets consisting of different data types—e.g. morphological, nucleotide, and protein—and to explore a wide variety of structured models mixing partition-unique and shared parameters. The program employs MPI to parallelize Metropolis coupling on Macintosh or UNIX clusters.
Availability: http://morphbank.ebc.uu.se/mrbayes
Contact: fredrik.ronquist@ebc.uu.se
*
To whom correspondence should be addressed.
The chloroplast DNA (cpDNA) inversion in the Asteraceae has been cited as a classic example of using genomic rearrangements for defining major lineages of plants. We further characterize cpDNA inversions in the Asteraceae using extensive sequence comparisons among 56 species, including representatives of all major clades of the family and related families. We determine the boundaries of the 22-kb (now known as 22.8 kb) inversion that defines a major split within the Asteraceae, and in the process, we characterize the second and a new, smaller 3.3-kb inversion that occurs at one end of the larger inversion. One end point of the smaller inversion is upstream of the trnE-UUC gene, and the other end point is located between the trnC-GCA and rpoB genes. Although a diverse sampling of Asteraceae experienced substantial length variation and base substitution during the long evolutionary history subsequent to the inversion events, the precise locations of the inversion end points are identified using comparative sequence alignments in the inversion regions. The phylogenetic distribution of two inversions is identical among the members of Asteraceae, suggesting that the inversion events likely occurred simultaneously or within a short time period shortly after the origin of the family. Estimates of divergence times based on ndhF and rbcL sequences suggest that two inversions originated during the late Eocene (38-42 MYA). The divergence time estimates also suggest that the Asteraceae originated in the mid Eocene (42-47 MYA).
We determined the distribution of a chloroplast DNA inversion among 80 species representing 16 tribes of the Asteraceae and 10 putatively related families. Filter hybridizations using cloned chloroplast DNA restriction fragments of lettuce and petunia revealed that this 22-kilobase-pair inversion is shared by 57 genera, representing all tribes of the Asteraceae, but is absent from the subtribe Barnadesiinae of the tribe Mutisieae, as well as from all families allied to the Asteraceae. The inversion thus defines an ancient evolutionary split within the family and suggests that the Barnadesiinae represents the most primitive lineage in the Asteraceae. These results also indicate that the tribe Mutisieae is not monophyletic, since any common ancestor to its four subtribes is also shared by other tribes in the family. This is the most extensive survey of the systematic distribution of an organelle DNA rearrangement and demonstrates the potential of such mutations for resolving phylogenetic relationships at higher taxonomic levels.
The evolutionary analysis of molecular sequence variation is a statistical enterprise. This is reflected in the increased use of probabilistic models for phylogenetic inference, multiple sequence alignment, and molecular population genetics. Here we present BEAST: a fast, flexible software architecture for Bayesian analysis of molecular sequences related by an evolutionary tree. A large number of popular stochastic models of sequence evolution are provided and tree-based models suitable for both within- and between-species sequence data are implemented.
BEAST version 1.4.6 consists of 81000 lines of Java source code, 779 classes and 81 packages. It provides models for DNA and protein sequence evolution, highly parametric coalescent analysis, relaxed clock phylogenetics, non-contemporaneous sequence data, statistical alignment and a wide range of options for prior distributions. BEAST source code is object-oriented, modular in design and freely available at http://beast-mcmc.googlecode.com/ under the GNU LGPL license.
BEAST is a powerful and flexible evolutionary analysis package for molecular sequence variation. It also provides a resource for the further development of new models and statistical methods of evolutionary analysis.
Evolutionary shifts to bird pollination (ornithophily) have occurred independently in many lineages of flowering plants. This shift affects many floral features, particularly those responsible for the attraction of birds, deterrence of illegitimate flower visitors (particularly bees), protection from vigorous foraging by birds, and accurate placement of pollen on bird's bodies. Red coloration appears to play a major role in both bee-deterrence and bird-attraction. Other mechanisms of bird-attraction include the production of abundant dilute nectar and the provision of secondary perches (for non-hovering birds). As a result of selection for similar phenotypic traits in unrelated bird-pollinated species, a floral syndrome of ornithophily can be recognized, and this review surveys the component floral traits. The strong convergent evolution evident in bird-pollinated flowers raises a question about the nature of the genetic mechanisms underlying such transitions and whether the same gene systems are involved in most cases. As yet there is too little information to answer this question. However, some promising model systems have been developed that include closely related bee and bird-pollinated flowers, such as Ipomoea, Mimulus, and Lotus. Recent studies of floral developmental genetics have identified numerous genes important in the development of the floral phenotype, which are also potential candidates for involvement in shifts between bee-pollination and bird pollination. As more whole-genome information becomes available, progress should be rapid.
Families of the Campanulales-Asterales-complex are characterized by special mechanisms for secondary pollen presentation: pump mechanism, brushing mechanism, deposition mechanism, cup mechanism and combination of cup mechanism with brushing mechanism. These different mechanisms are based on three successive events: elongation of the filaments, opening of the anthers, elongation of the style. The diversity of the mechanisms arises through different auxiliary structures such as an indusium, hairs or an anther tube, and the relative rate and time of filament and style growth. An advantage of secondary pollen presentation lies in the prolongation of the male phase of anthesis through portioned pollen release. A probable phylogeny of the different mechanisms of secondary pollen presentation is proposed (Fig. 7).
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
The effects of different Nb and Fe addition ratios on the microstructure, corrosion and oxide characteristics of Zr-based alloys were investigated. The Nb/Fe ratio was controlled to be 0.6, 1.0, 1.7, 3.0, and 7.0 with the same amount of Nb+Fe in each alloy. The microstructural analysis and precipitate characterization were performed to obtain the correlation between the corrosion and the microstructures. The grain size and the area fraction of the precipitate in all the tested alloys were almost the same even though the Nb/Fe ratio was considerably changed. But the mean diameter and the crystal structure of the precipitates were affected by the variation of the Nb/Fe ratio. In the alloy with a low Nb content, the FCC-(ZrNb)2Fe precipitate was mainly formed while the HCP-Zr(NbFe)2 precipitate was frequently observed in the alloy of with a high Nb content. The corrosion resistance of the Zr–xNb–yFe ternary alloys was improved by decreasing the Nb/Fe ratio. From the microstructure and corrosion studies, it seems that the corrosion resistance would be more closely related to the crystal structure of the precipitate rather than the other properties such as the size distribution and the density of the precipitates.
The systematic position of Feddea (Asteraceae) has been enigmatic ever since its publication in 1925. The latest taxonomic accounts of the family left it either unplaced within subfamily Asteroideae or unplaced within tribe Inuleae. Macro-morphological and electron microscopy data (scanning electron microscopy and transmission electron microscopy) indicated that Feddea is not part of an early branching lineage of the Asteraceae, contrary to what had been suggested in initial taxonomic accounts. Feddea has style branches with a 2-banded (vs. continuous) stigmatic surface, thus differing from early diverging lineages of the family. Among Asteroideae, Feddea is diagnosed by discoid capitula with all florets bisexual, long-caudate anthers, noncarbonized cypselae, and pollen with a narrow cavus region, columellae without internal foramina, but with a complex basal columellate layer. A phylogenetic reconstruction based on nucleotide sequences of the chloroplast gene ndhF showed that Feddea is sister to the Heliantheae s.l. There were, however, no clear morphological synapomorphies shared with that tribe and we therefore propose Feddeeae as a new unispecific tribe to accommodate this critically endangered genus restricted to Eastern Cuba.
Abstract New inference techniques, such as supertrees, have improved the construction of large phylogenies, helping to reveal the tree of life. In addition, these large phylogenies have enhanced the study of other evolutionary questions, such as whether traits have evolved in a neutral or adaptive way, or what factors have influenced diversification. However, supertrees usually lack branch lengths, which are necessary for all these issues to be investigated. Here, divergence times within the largest family of flowering plants, namely the Asteraceae, are reviewed to estimate time-calibrated branch lengths in the supertree of this lineage. An inconsistency between estimated dates of basal branching events and the earliest asteraceous fossil pollen record was detected. In addition, the impact of different methods of branch length assignment on the total number of transitions between states in the reconstruction of sexual system evolution in Asteraceae was investigated. At least for this dataset, different branch length assignation approaches influenced maximum likelihood (ML) reconstructions only and not Bayesian ones. Therefore, the selection of different branch length information is not arbitrary and should be carefully assessed, at least when ML approaches are being used. The reviewed divergence times and the estimated time-calibrated branch lengths provide a useful tool for future phylogenetic comparative and macroevolutionary studies of Asteraceae.
Not available Biological Sciences, School of
We describe MUSCLE, a new computer program for creating multiple alignments of protein sequences. Elements of the algorithm
include fast distance estimation using kmer counting, progressive alignment using a new profile function we call the log‐expectation score, and refinement using tree‐dependent
restricted partitioning. The speed and accuracy of MUSCLE are compared with T‐Coffee, MAFFT and CLUSTALW on four test sets
of reference alignments: BAliBASE, SABmark, SMART and a new benchmark, PREFAB. MUSCLE achieves the highest, or joint highest,
rank in accuracy on each of these sets. Without refinement, MUSCLE achieves average accuracy statistically indistinguishable
from T‐Coffee and MAFFT, and is the fastest of the tested methods for large numbers of sequences, aligning 5000 sequences
of average length 350 in 7 min on a current desktop computer. The MUSCLE program, source code and PREFAB test data are freely
available at http://www.drive5. com/muscle.
The darwinian concept of "descent with modification" applies to metabolic pathways: pathways sharing similarities must have inherited them from an exclusive, hypothetical ancestral pathway. Comparative anatomy of biochemical pathways is performed using five criteria of homology. Primary homologies of "type I" were defined as several pathways sharing the same enzyme with high specificity for its substrate. Primary homologies of "type II" were defined as the sharing of similar enzymatic functions, cofactors, functional family, or recurrence of a set of reactions. Standard cladistic analysis is used to infer the evolutionary history of metabolic development and the relative ordering of biochemical reactions through time, from a single matrix integrating the whole basic universal metabolism. The cladogram shows that the earliest pathways to emerge are metabolism of amino acids of groups I and II (Asp, Asn, Glu, and Gln). The earliest enzymatic functions are mostly linked to amino acid catabolism: deamination, transamination, and decarboxylation. For some amino acids, catabolism and biosynthesis occur at the same time (Asp, Glu, Lys, and Met). Catabolism precedes anabolism for Asn, Gln, Arg, Trp, His, Tyr, and Phe, and anabolism precedes catabolism for Pro, Ala, Leu, Val, Ile, Cys, Gly, Ser, and Thr. The urea cycle evolves from arginine synthesis. Metabolism of fatty acids and sugars develops after the full development of metabolism of amino acids of groups I and II, and they are associated with the anabolism of amino acids of groups III and IV. Syntheses of aromatic amino acids are branched within sugar metabolism. The Krebs cycle occurs relatively late after the setting of metabolism of amino acids of groups I and II. One portion of the Krebs cycle has a catabolic origin, whereas the other portion has an anabolic origin in pathways of amino acids of groups III and IV. It is not possible to order glycolysis and gluconeogenesis with regard to the Krebs cycle, as they all belong to "period 6." Pentose-phosphate and Calvin cycles are later (periods 7 and 8, respectively). Cladistic analysis of the structure of biochemical pathways makes hypotheses in biochemical evolution explicit and parsimonious.
The largest family of flowering plants Asteraceae (Compositae) is found to contain 12 major lineages rather than five as previously suggested. Five of these lineages heretofore had been circumscribed in tribe Mutisieae (Cichorioideae), a taxon shown by earlier molecular studies to be paraphyletic and to include some of the deepest divergences of the family. Combined analyses of 10 chloroplast DNA loci by different phylogenetic methods yielded highly congruent well-resolved trees with 95% of the branches receiving moderate to strong statistical support. Our strategy of sampling genera identified by morphological studies as anomalous, supported by broader character sampling than previous studies, resulted in identification of several novel clades. The generic compositions of subfamilies Carduoideae, Gochnatioideae, Hecastocleidoideae, Mutisioideae, Pertyoideae, Stifftioideae, and Wunderlichioideae are novel in Asteraceae systematics and the taxonomy of the family has been revised to reflect only monophyletic groups. Our results contradict earlier hypotheses that early divergences in the family took place on and spread from the Guayana Highlands (Pantepui Province of northern South America) and raise new hypotheses about how Asteraceae dispersed out of the continent of their origin. Several nodes of this new phylogeny illustrate the vast differential in success of sister lineages suggesting focal points for future study of species diversification. Our results also provide a backbone exemplar of Asteraceae for supertree construction.
Despite extensive efforts, parts of the phylogeny of the angiosperm family Rubiaceae has not been resolved and consequently, character evolution, ancestral areas and divergence times of major radiations are difficult to estimate. Here, phylogenetic analyses of 149 taxa and five plastid gene regions show that three enigmatic genera are sisters to considerably species rich clades. The rare and endangered species Dunnia, endemic to southern Guangdong, China, is sister to a large clade in the Spermacoceae alliance; the rarely collected Schizocolea from western tropical Africa is sister to the Psychotrieae alliance; and Colletoecema from central tropical Africa is sister to remaining Rubioideae. The morphology of these taxa has been considered "puzzling". In combination with further morphological studies, our results may help understanding the apparently confusing traits of these plants. Phylogenetic, morphological, and geographical isolation of Dunnia, Schizocolea and Colletocema may indicate high genetic diversity. They are lone representatives of unique lineages and if extinct, the loss would not only mean loss of genetic diversity of a single species but of an entire lineage.
An RNA secondary structure model is presented for the nuclear ribosomal internal transcribed spacers (ITS) based on comparative analysis of 340 sequences from the angiosperm family Asteraceae. The model based on covariation analysis agrees with structural features proposed in previous studies using mainly thermodynamic criteria and provides evidence for additional structural motifs within ITS1 and ITS2. The minimum structure model suggests that at least 20% of ITS1 and 38% of ITS2 nucleotide positions are involved in base pairing to form helices. The sequence alignment enabled by conserved structural features provides a framework for broadscale molecular evolutionary studies and the first family-level phylogeny of the Asteraceae based on nuclear DNA data. The phylogeny based on ITS sequence data is very well resolved and shows considerable congruence with relationships among major lineages of the family suggested by chloroplast DNA studies, including a monophyletic subfamily Asteroideae and a paraphyletic subfamily Cichorioideae. Combined analyses of ndhF and ITS sequences provide additional resolution and support for relationships in the family.
The history of the development of statistical hypothesis testing in time series analysis is reviewed briefly and it is pointed out that the hypothesis testing procedure is not adequately defined as the procedure for statistical model identification. The classical maximum likelihood estimation procedure is reviewed and a new estimate minimum information theoretical criterion (AIC) estimate (MAICE) which is designed for the purpose of statistical identification is introduced. When there are several competing models the MAICE is defined by the model and the maximum likelihood estimates of the parameters which give the minimum of AIC defined by AIC = (-2)log-(maximum likelihood) + 2(number of independently adjusted parameters within the model). MAICE provides a versatile procedure for statistical model identification which is free from the ambiguities inherent in the application of conventional hypothesis testing procedure. The practical utility of MAICE in time series analysis is demonstrated with some numerical examples.
Arctotis hirsuta (Harv.) P. Beauv., Panero 2002-61, Culti-vated, seed source: Kirstenboch Botanical Garden Artemisia frigida Willd Chuqui-raga spinosa Less
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, KM191983, EU385317, EU385220, EU385125, EU243243, KM191818, EU384939, EU385411, EU385507, KM191922, EU385031. Ainsliaea apiculata Sch. Bip ex. Zoll., Japan, no voucher. KM192036, KM191984, EU385321, EU385225, EU385130, EU243248, KM191819, EU384944, EU385416, EU385512, KM191923, EU385036. Aphyllocladus spartioides Wedd., Panero and Crozier 8500, Argentina, SI, TEX. KM192037, KM191985, EU385323, EU385227, EU385132, EU243250, KM191820, EU384946, EU385418, EU385513, KM191924, EU385038. Arctotis hirsuta (Harv.) P. Beauv., Panero 2002-61, Culti-vated, seed source: Kirstenboch Botanical Garden, South Africa, TEX. KM192038, KM191986, EU385324, EU385228, EU385133, EU243251, KM191821, EU384947, EU385419, EU385514, KM191925, EU385039. Artemisia frigida Willd., NM1, China, no voucher. NC020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607, NC 020607. Athroisma gracile (Oliv.) Mattf. ssp. psyllioides (Oliv.) T. Eriksson, Eriksson, Kalema, and Leliyo 559, Tanzania, TEX. AY214947, KJ434416, AY215765, AF384437, L39455, AF383757, AY217218, AY215085, AY213763, EU385515, KM191926, AY216019/AY216144. Atractylodes japonica Koidz., cul-tivated Asiatica Nursery, no voucher, TEX. KM192039, KM191987, KM192111, KM191890, KM192100, KM191868, KM191822, KM192089, KM191879, KM191912, KM191927, KM191901. Barna-desia odorata Griseb., Panero and Crozier 8492, Argentina, TEX. KM192040, KM191988, EU385327, EU385231, NS, EU243254, KM191823, EU841102, EU385422, EU385518, KM191928, EU345042. Boopis anthemoides Juss., Simon 258, Argentina, WU. NS, NS, EU841363, NS, NS, NS, NS, EU841136, NS, NS, NS, EU841095/EU547627. Brachylaena elliptica (Thunb.) DC., Koekem-oer and Funk 1971, South Africa, US. KM192041, KM191989, EU385330, EU385234, EU385138, EU243357, KM191824, EU384952, EU385425, EU385521, KM191929, EU385045. Calopap-pus acerosus Meyen, Panero and Crozier 8457, Chile, CONC, TEX. KM192042, KM191990, KM192112, KM191891, KM192101, KM191869, KM191825, KM192090, KM191880, KM191913, KM191930, KM191902. Calycera crassifolia Hicken., Panero and Crozier 8392, Argentina, SI, TEX. KM192043, KM191991, KM192113, KM191892, KM192102, KM191870, KM191826, KM192091, KM191881, KM191914, KM191931, KM191903. Chap-talia nutans (L.) Pol., Panero 2002-19, USA, TEX. KM192044, KM191992, EU385335, EU385239, EU385143, EU243262, KM191827, EU384957, EU385430, EU385526, KM191932, EU385050. Chimantaea humilis Maguire, Steyermark & Wurdack, Weitzman et al. 412, Venezuela, US. KM192045, KM191993, EU385336, EU385240, EU385144, EU243263, KM191828, EU384958, EU385431, EU385527, KM191933, EU385051. Chuqui-raga spinosa Less., Simon 522, Argentina, US. KM192046, KM191994, EU385338, EU385242, EU385146, EU243265, KM191829, EU384960, EU385433, EU385529, KM191934, EU385053. Cnicothamnus lorentzii Griseb., Panero 1934, Argentina, TENN. KM192047, KM191995, EU385339, EU385243, EU385147, EU243266, KM191830, EU384961, EU385434, EU385530, KM191935, EU385054. Coreopsis petrophiloides B. L. Rob. & Greenm., Panero 3047, Mexico, TEX. AY214958, KJ434420, AY215776, AF384448, AF384704, AF383768, AY217229, AY215096, AY213774, AY213774, KM191936, AY216030/ AY216155. Corymbium glabrum L., Moffett 8764, South Africa, US. KM192048, KM191996, EU385340, EU385344, EU385148, EU243267, KM191831, EU384962, EU385435, EU385531, KM191937, EU385055. Cyclolepis genistoides D. Don, Panero and Crozier 8387, Argentina, SI, TEX. KM192049, KM191997, EU385341, EU385345, EU385149, EU243268, KM191832, EU384963, EU385436, EU385532, KM191938, EU385056. Dasyphyllum reticulatum (DC.) Cabrera, Roque, Funk and Kim 485, Brazil, US. KM192050, KM191998, EU385342, EU385246, EU385150, EU243269, KM191833, AY874428, EU385437, EU385533, KM191939, EU385057. Doniophyton anomalum (D. Don) Kurtz, Bonifacino 96, Argentina, US. KM192052, KM192000, EU385348, EU385252, EU385156, EU243274, KM191835, EU384969, EU385443, EU385538, KM191941, EU385063. Duidaea pinifolia S.F. Blake, V.A. Funk 8010, Venezuela, US. KM192053, KM192001, EU385349, EU385253, EU385157, EU243275, KM191836, EU384970, EU385444, EU385539, KM191942, EU385064. Duseniella patagonica (O. Hoffm.) K. Schum., Urtubey 149-2006, Argentina, LP. NS, NS, NS, NS, NS, NS, NS, NS, NS, NS, NS, EU547652. Echinops ritro L., Panero 2002-71. Cultivated, TEX. KM192054, KM192002, EU385350, EU385254, EU385158, EU243276, KM191837, EU384971, EU385445, EU385540, KM191943, EU385065. Famatinanthus decussatus Ariza and S.E.
Mutisia retrorsa Cav Nastanthus caespitosus (Phil Perezia recurvata Less
- Cultivated
- South Garden
- Tex Africa
- Km
- Km
- Eu
- Eu
- Eu
- Eu
- Km
- Eu
- Eu
- Eu
- Eu385090 Km
- Marshallia Caespitosa Nutt
- Ex
- Dc
- Panero
- Tex Usa
- Ay
- Kj
- Ay
- Af
- L
- Af
- Ay
- Ay
- Ay
- Ay
- Ay216077 Km
- Ay216202 Menyanthes
- Tex Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km191908 Km
- Us Argentina
- Km
- Ns
- Eu
- Eu
- Eu
- Eu
- Ns
- Eu
- Eu
- Eu
- Eu385091 Ns
- Chile
- Tex Conc
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km191909 Km
- Nouelia
- Franch
- Us China
- Km
- Km
- Eu
- Eu
- Eu
- Eu
- Km
- Eu
- Eu
- J L Eu
- Panero
Cultivated, seed source: Kirstenbosch Botanical Garden, South Africa, TEX. KM192071, KM192019, EU385375, EU385280, EU385184, EU243302, KM191853, EU384997, EU385471, EU385568, KM191963, EU385090. Marshallia caespitosa Nutt. Ex. DC., Panero 7426, USA, TEX. AY215006, KJ434430, AY215824, AF384496, L39458, AF383816, AY217274, AY215142, AY213820, AY213820, KM191964, AY216077/AY216202. Menyanthes trifoliata L., Panero 2011-1, cultivated, TEX. KM192072, KM192020, KM192118, KM191897, KM192107, KM191875, KM191854, KM192096, KM191886, KM191918, KM191965, KM191908. Mutisia retrorsa Cav., Bonifacino 148, Argentina, US. KM192073, NS, EU385376, EU385281, EU385185, EU243303, NS, EU384998, EU385472, EU385569, NS, EU385091. Nastanthus caespitosus (Phil.) Reiche, Panero and Crozier 8431, Chile, CONC, TEX. KM192074, KM192021, KM192119, KM191898, KM192108, KM191876, KM191855, KM192097, KM191887, KM191919, KM191966, KM191909. Nouelia insignis Franch., Rock 8534, China, US. KM192075, KM192022, EU385378, EU385283, EU385187, EU243305, KM191856, EU385000, EU385474, EU385571, J.L. Panero et al. / Molecular Phylogenetics and Evolution 80 (2014) 43–53 KM191967, EU385093. Oldenburgia grandis (Thunb.) Baill., B & T World Seeds, South Africa, no voucher. KM192076, KM192023, EU385379, EU385284, EU385188, EU243306, KM191857, EU385001, EU385475, EU385572, KM191968, EU385094. Onoseris hastata Wedd., cultivated, Fairchild Gardens, no voucher. KM192077, KM192024, EU385381, EU385286, EU385190, EU243308, KM191858, EU385003, EU385477, EU385574, KM191969, EU385096. Pachylaena atriplicifolia D. Don ex H. & A., Panero and Crozier 8477A, Argentina, SI, TEX. KM192078, KM192025, EU385383, EU385288, EU385192, EU243310, KM191859, EU385005, EU385479, EU385577, KM191970, EU385098. Perezia recurvata Less., Panero and Crozier 8399, Argentina, SI, TEX. KM192079, KM192026, KM192120, KM191899, KM192109, KM191877, KM191860, KM192098, KM191888, KM191920, KM191971, KM191910. Pertya scandens Sch. Bip., Japan, no voucher. KM192080, KM192027, EU385386, EU385291, EU385195, EU243313, KM191861, EU385008, EU385482, EU385580, KM191972, EU385101. Phaneroglossa bolusii (Oliv.) B. Nordenstam, Watson and Panero 94-62, South Africa, TEX. AY215025, KM192028, AY215843, AF384514, AF384765, AF383835, AY217293, AY215161, AY213836, AY213836, KM191973, AY216096/AY216221. Plazia daphnoides Wedd., Panero and Crozier 8439, Chile, CONC, TEX. KM192081, KM192029, EU385388, EU385293, EU385197, EU243315, KM191862, EU385010, EU385484, EU385582, KM191974, EU385103. Schlechtendalia luzulifolia Less., Panero and Crozier 8535, Argentina, TEX, SI. KM192082, KM192030, KM192121, KM191900, KM192110, KM191878, KM191863, KM192099, KM191889, KM191921, KM191975, KM191911. Sinclairia palmeri (A. Gray) B.L. Turner, Panero 7457, Mexico, TEX. KM192068, KM192016, EU385373, EU385277, EU385181, EU243299, KM191850, EU384994, EU385468, EU385565, KM191960, EU385087. Stenopadus talaumifolius S. F. Blake, Clarke 5459, Venezuela, US. KM192083, NS, EU385398, EU385303, EU385207, EU243323, NS, EU385019, EU385494, EU385591, KM191976, EU385113. Stifftia chrysantha Mikan, Serra 235, Brazil, TEX. KM192084, KM192031, EU385399, EU385304, EU385208, EU243324, KM191864, EU385020, EU385495, EU385592, KM191977, EU385114. Stomatochaeta condensata (Baker) Maguire & Wurdack, Berry 6574B, Venezuela, US. KM192085, NS, EU385401, EU385306, EU385210, EU243326, NS, EU385021, EU385497, EU385594, KM191978, EU385115. Tagetes erecta L., Soule 3004, Guatemala, TEX. AY215049, KJ434436, AY215867, AF384536, L39466, AF383858, AY217315, AY215184, AY213858, AY213858, KM191979, AY216119/AY216244. Trixis divaricata (Kunth) Spreng., Santos 2659, Brazil, TEX. KM192086, KM192032, EU385405, EU385310, EU385214, EU243330, KM191865, EU385025, EU385501, EU385598, KM191980, EU385120. Wunderlichia mirabi-lis Riedel, Roque, Funk and Kim 466, Brazil, US. KM192087, KM192033, EU385408, EU385313, EU385217, EU243333, KM191866, EU385028, EU385504, EU385601, KM191981, EU385122. Youngia japonica (L.) DC., Panero 2002-92, USA, TEX. KM192088, KM192034, EU385409, EU385314, EU385218, EU243334, KM191867, EU385029, EU385505, EU385602, KM191982, EU385123. Appendix B. Supplementary material Supplementary data associated with this article can be found, in the online version, at http://dx.doi.org/10.1016/j.ympev.2014.07. 012. References Akaike, H., 1974. A new look at the statistical model identification. IEEE Trans. Automat. Control 19, 716–723.
Hyaloseris salicifolia (Griseb.) Hieron., Simon 330
- Cultivated
- South Garden
- Tex Africa
- Km
- Km
- Eu
- Eu
- Eu
- Eu
- Km
- Eu
- Eu
- Eu
- Eu385068 Km
- H V Gamocarpha
- Hansen
- Crozier Panero
- Chile
- Tex Conc
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km191905 Km
- ( Dc. ) A Gochnatia
- Gray
- Mex-1 Panero
- Tex Mexico
- Km
- Km
- Eu
- Eu
- Eu
- Eu
- Km
- Eu
- Eu
- Eu
- Eu385071 Km
- R H Gongylolepis
- Schomb
- Berry
- Us Venezuela
- Km
- Ns
- Eu
- Eu
- Eu
- Eu
- Ns
- Eu
- Eu
- Eu
- Eu385073 Km
- Gymnarrhena
- Desf
- Saudi Mandeville
- Us Arabia
- Km
- Km
- Eu
- Eu
- Eu
- Eu
- Km
- Eu
- Eu
- Eu
- Eu385076 Km
- Chile
- Tex Conc
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km
- Km191906 Km
- A Hecastocleis
- Gray
- Crozier Panero
- Tex Usa
- Km
- Km
- Eu
- Eu
- Eu
- Eu
- Km
- Eu
- Eu
- Eu
- Eu385077 Km
- L Helianthus
- Tex Ay
- Kj
- Ay
- Af
- L
- Af
- Ay
- Ay
- Ay
- Eu
- Ay216058 Km
- D J N Ay
- Hind
Cultivated, seed source: Kirstenbosch Botanical Garden, South Africa, TEX. KM192056, KM192004, EU385353, EU385257, EU385161, EU243279, KM191839, EU384974, EU385448, EU385543, KM191945, EU385068. Gamocarpha alpina (Poepp. ex Less.) H.V. Hansen, Panero and Crozier 8414, Chile, CONC, TEX. KM192057, KM192005, KM192115, KM191894, KM192104, KM191872, KM191840, KM192093, KM191883, KM191915, KM191946, KM191905. Gochnatia hypoleuca (DC.) A. Gray, Panero MEX-1, Mexico, TEX. KM192058, KM192006, EU385357, EU385261, EU385165, EU243283, KM191841, EU384978, EU385452, EU385546, KM191947, EU385071. Gongylolepis benthamiana R. H. Schomb., Berry 6564, Venezuela, US. KM192059, NS, EU385359, EU385263, EU385167, EU243285, NS, EU384980, EU385454, EU385547, KM191948, EU385073. Gymnarrhena micrantha Desf., Mandeville 157, Saudi Arabia, US. KM192060, KM192007, EU385362, EU385266, EU385170, EU243288, KM191842, EU384983, EU385457, EU385550, KM191949, EU385076. Gypothamnium pinifolium Phil., Panero and Crozier 8437, Chile, CONC, TEX. KM192061, KM192008, KM192116, KM191895, KM192105, KM191873, KM191843, KM192094, KM191884, KM191916, KM191950, KM191906. Hecastocleis shockleyi A. Gray, Panero and Crozier 8157, USA, TEX. KM192062, KM192009, EU385363, EU385267, EU385171, EU243289, KM191844, EU384984, EU385458, EU385551, KM191951, EU385077. Helianthus annuus L., Panero 2002-27, USA, TEX. AY214986, KJ434427, AY215805, AF384476, L39383, AF383796, AY217256, AY215124, AY213802, EU385553, KM191952, AY216058/AY216183. Hesperomannia arbuscula Hillebr., cultivated, University of Hawaii, USA, no vou-cher. KM192063, KM192010, EU385364, EU385268, EU385172, EU243290, KM191845, EU384985, EU385459, EU385554, KM191953, EU385078. Hofmeisteria fasciculata Brandegee, Panero 2817, Mexico, TEX. AY214987, KM192011, AY215806, AF384477, AF384731, AF383797, AY217257, AY215125, AY213803, AY213803, KM191954, AY216059/AY216184. Huarpea andina Cabrera, Dalmasso s.n., Argentina, LP. NS, NS, EU841349, NS, NS, NS, NS, EU841124, NS, NS, NS, EU841084/EU547653. Hyalis argentea D. Don ex Hook. & Arn., Panero and Crozier 8388, Argentina, TEX. KM192064, KM192012, EU385367, EU385271, EU385175, EU243293, KM191846, EU384988, EU385462, EU385557, KM191955, EU385081. Hyaloseris salicifolia (Griseb.) Hieron., Simon 330, Argentina, US. KM192051, KM191999, EU385346, EU385250, EU385154, EU243272, KM191834, EU384967, EU385441, EU385536, KM191940, EU385061. Hyaloseris rubicunda Griseb., Simon 716, Argentina, US. KM192065, KM192013, EU385368, EU385272, EU385176, EU243294, KM191847, EU384989, EU385463, EU385558, KM191956, EU385082. Ianthopappus corymbosus (Less.) Roque & D.J.N. Hind, Panero and Crozier 8530, Argentina, SI, TEX. KM192066, KM192014, EU385369, EU385273, EU385177, EU243295, KM191848, EU384990, EU385464, EU385559, KM191957, EU385083. Inula britannica L., Santos and Francisco ACC55-98, Cultivated, ORT. AY214993, KJ434428, AY215812, AF384483, AF384737, AF383803, AY217263, AY215130, AY213809, AY213809, KM191958, AY216065/AY216190. Leucome-ris spectabilis D. Don, Nicolson 3254, Nepal, US. KM192067, KM192015, EU385372, EU385276, EU385180, EU243298, KM191849, EU384993, EU385467, EU385564, KM191959, EU385086. Lophopappus cuneatus R.E. Fr., Panero and Crozier 8508, Argentina, SI, TEX. KM192069, KM192017, EU385374, EU385278, EU385182, EU243300, KM191851, EU384995, EU385469, EU385566, KM191961, EU385088. Lycoseris latifolia (D. Don) Benth., Panero 2626, Venezuela, MY. KM192070, KM192018, KM192117, KM191896, KM192106, KM191874, KM191852, KM192095, KM191885, KM191917, KM191962, KM191907. Macledium zeyheri (Sond.) S. Ortiz, Panero 2002-47.
JModelTest 2: more models, new heuristics and parallel computing
- Darriba
Darriba, D., Taboada, G.L., Doallo, R., Posada, D., 2012. JModelTest 2: more models,
new heuristics and parallel computing. Nat. Methods 9, 772.
Families and Genera of Vascular Plants, Flowering Plants
- C Jeffrey
Jeffrey, C., 2007. Compositae: introduction with key to tribes. In: Kadereit, J.W.,
Jeffrey, C. (Eds.), Families and Genera of Vascular Plants, Flowering Plants,
Eudicots, Asterales, vol. VIII. Springer-Verlag, Berlin, pp. 61-87.
Two chloroplast DNA inversions originated simultaneously during the early evolution of the sunflower family
- H G Kim
- K S Choi
- R K Jansen
Kim, H.G., Choi, K.S., Jansen, R.K., 2005. Two chloroplast DNA inversions originated
simultaneously during the early evolution of the sunflower family (Asteraceae).
Portioned pollen release and the syndromes of secondary pollen presentation in the Campanulales–Asterales complex
- Erbar