The Paraphyly of Osmunda is Confirmed by Phylogenetic Analyses of Seven Plastid Loci (original) (raw)
Related papers
The Paraphyly of Osmunda is Confirmed by Phylogenetic Analyses of Seven Plastid Loci
Systematic Botany, 2008
To resolve phylogenetic relationships among all genera and subgenera in Osmundaceae, we analyzed over 8,500 characters of DNA sequence data from seven plastid loci (atpA, rbcL, rbcL-accD, rbcL-atpB, rps4-trnS, trnG-trnR, and trnL-trnF). Our results confirm those from earlier anatomical and single-gene (rbcL) studies that suggested Osmunda s.l. is paraphyletic. Osmunda cinnamomea is sister to the remainder of Osmundaceae (Leptopteris, Todea, and Osmunda s.s.). We support the recognition of a monotypic fourth genus, Osmundastrum, to reflect these results. We also resolve subgeneric relationships within Osmunda s.s. and find that subg. Claytosmunda is strongly supported as sister to the rest of Osmunda. A stable, well-supported classification for extant Osmundaceae is proposed, along with a key to all genera and subgenera.
Journal of Plant Research, 1999
We determined rbcL sequences of tl of 15 extant species of Osmundaceae which represent all three genera, Osmunda, Todea and Leptopterk Our phylogenetic analysis concluded: 1) Osmunda subg. Osmunda and subg. Pknasium are monophyletic groups, but subg. Osmundastrum is not. The genus Osmunda is not monophyletic because T W and Leptopteris are positioned within Osmunda. 2) Osmunda cinnamomea is the most basally positioned species in Osmundaceae, and it can be called as "a living fossil" because a fossil species (0. c/aytoniites) with almost the same morphology as this species was recorded from the Triassic. 3) Osmunda jbpor~ica and 0. regals are very closely related with only one nucleotii difference in the rbcL gene. 4) Greater nucleotide variation (5-7 nucleotides) was found between conspecific samples of 0. cinnamomea and 0. c/aytonbna collected from Japan and United States. Each of these two species may comprise more than two biologically differentiated species.
Phylogenetics of sees plants: An analysis of nucleotide sequences from the plastid gene rbcL
1993
We present the results of two exploratory parsimony analyses of DNA sequences from 475 and 499 species of seed plants, respectively, representing all major taxonomic groups. The data are exclusively from the chloroplast gene rbcL, which codes for the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO or RuBPCase). We used two different state-transformation assumptions resulting in two sets of cladograms: (i) equal-weighting for the 499-taxon analysis; and (ii) a procedure that differentially weights transversions over transitions within characters and codon positions among characters for the 475-taxon analysis. The degree of congruence between these results and other molecular, as well as morphological, cladistic studies indicates that rbcL sequence variation contains historical evidence appropriate for phylogenetic analysis at this taxonomic level of sampling. Because the topologies presented are necessarily approximate and cannot be evaluated adequately for internal support, these results should be assessed from the perspective of their predictive value and used to direct future studies, both molecular and morphological. In both analyses, the three genera of Gnetales are placed together as the sister group of the flowering plants, and the anomalous aquatic Ceratophyllum (Ceratophyllaceae) is sister to all other flowering plants. Several major lineages identified correspond well with at least some recent taxonomic schemes for angiosperms, particularly those of Dahlgren and Thorne. The basalmost clades within the angiosperms are orders of the apparently polyphyletic subclass Magnoliidae sensu Cronquist. The most conspicuous feature of the topology is that the major division is not monocot versus dicot, but rather one correlated with general pollen type: uniaperturate versus triaperturate. The Dilleniidae and Hamamelidae are the only subclasses that are grossly polyphyletic; an examination of the latter is presented as an example of the use of these broad analyses to focus more restricted studies. A broadly circumscribed Rosidae is paraphyletic to Asteridae and Dilleniidae. Subclass Caryophyllidae is monophyletic and derived from within Rosidae in the 475-taxon analysis but is sister to a group composed of broadly delineated Asteridae and Rosidae in the 499-taxon study.
Background: The classification of royal ferns (Osmundaceae) has long remained controversial. Recent molecular phylogenies indicate that Osmunda is paraphyletic and needs to be separated into Osmundastrum and Osmunda s.str. Here, however, we describe an exquisitely preserved Jurassic Osmunda rhizome (O. pulchella sp. nov.) that combines diagnostic features of both Osmundastrum and Osmunda, calling molecular evidence for paraphyly into question. We assembled a new morphological matrix based on rhizome anatomy, and used network analyses to establish phylogenetic relationships between fossil and extant members of modern Osmundaceae. We re-analysed the original molecular data to evaluate root-placement support. Finally, we integrated morphological and molecular data-sets using the evolutionary placement algorithm. Results: Osmunda pulchella and five additional Jurassic rhizome species show anatomical character suites intermediate between Osmundastrum and Osmunda. Molecular evidence for paraphyly is ambiguous: a previously unrecognized signal from spacer sequences favours an alternative root placement that would resolve Osmunda s.l. as monophyletic. Our evolutionary placement analysis identifies fossil species as probable ancestral members of modern genera and subgenera, which accords with recent evidence from Bayesian dating. Conclusions: Osmunda pulchella is likely a precursor of the Osmundastrum lineage. The recently proposed root placement in Osmundaceae—based solely on molecular data—stems from possibly misinformative outgroup signals in rbcL and atpA genes. We conclude that the seemingly conflicting evidence from morphological, anatomical, molecular, and palaeontological data can instead be elegantly reconciled under the assumption that Osmunda is indeed monophyletic.
Botanical Journal of the Linnean Society
Phylogenetic and molecular clock analyses were performed including all genera except one (Pseudomonotes) for the three subfamilies of Dipterocarpaceae. We also included representatives of Sarcolaenaceae and Cistaceae with Bixaceae as the ultimate outgroup. Three plastid regions (six markers), partial rbcL, trnK-matK-trnK (partial trnK intron including complete matK) and trnT-trnL-trnF (partial trnT, complete trnT-trnL intergenic spacer, complete trnL, complete trnL-trnF intergenic spacer and partial trnF), were analysed. We also investigated additional accessions for genome size and chromosome numbers. Our phylogenetic results differ in three important respects from previous interpretations of morphological characters, as reflected in recent classifications. First, our analyses strongly support assignment of Pakaraimaea (subfamily Pakaraimaeoideae) to Cistaceae. Second, the morphological concepts of Dipterocarpeae and Shoreeae in subfamily Dipterocarpoideae are not supported because Dipterocarpus is sister to Dryobalanops plus tribe Shoreeae. Our analysis revealed four clades: (1) Dipterocarpus; (2) Dryobalanops, for which tribal assignment has been contentious; (3) genera of Shoreeae; and (4) the remaining genera of Dipterocarpeae. Third, Shorea is not monophyletic. Monotoideae are weakly supported as sister to Dipterocarpoideae; Sarcolaenaceae (endemic to Madagascar) are sister to this pair. Divergence in extant Dipterocarpoideae occurred c. 55 Mya. Genome sizes for all accessions examined are small (0.3264-0.6724 pg), and the additional chromosome numbers we collected fit into the patterns previously observed for Dipterocarpaceae.
Molecular phylogenetics of Linaceae with complete generic sampling and data from two plastid genes
Botanical Journal of the Linnean Society, 2011
The phylogeny of Linaceae is examined, with sampling from the 13 commonly recognized genera of the family and sequence data from the plastid genes matK and rbcL. Representatives of 24 additional families of the order Malpighiales are included in the analyses, with members of Celastrales, Fabales, Fagales, Oxalidales and Rosales used as outgroups. Linaceae and both subfamilies, the temperate Linoideae and the tropical Hugonioideae, are found to be monophyletic in likelihood-and parsimony-based analyses, although the monophyly of Hugonioideae is not well supported. Average divergence time estimates using rbcL indicate that the subfamilies diverged from each other during the Palaeocene, approximately 60 million years ago. No sister group to Linaceae is consistently identified in these analyses, and relationships among families of Malpighiales are not well resolved. In accord with previous estimates of Linoideae phylogeny, Linum is shown to be nonmonophyletic, with several segregate genera nested within it, but the relationships of the south-east Asian genera, Anisadenia, Reinwardtia and Tirpitzia, remain uncertain. In Hugonioideae, Indorouchera and Philbornea are found to be closely related to members of Hugonia section Durandea. Relationships of the neotropical genera Hebepetalum and Roucheria to the palaeotropical hugonioids are not consistently resolved.
Botanical Journal of The Linnean Society, 2010
Phylogenetic relationships within the palaeotropical tribe Lepisoroideae (Polypodiaceae) were investigated by studying sequence variation of four plastid DNA regions: rbcL, rps4 plus rps4-trnS IGS, trnL intron plus trnL-F IGS, rbcL-atpB IGS plus part of atpB. In total, over 4000 nucleotides were sequenced for 39 species. Seven well-supported clades were found in the analyses of the combined data set. We provide a new classification of Lepisoroideae by integrating phylogenetic results and known variation of morphological characters. The two small genera Neocheiropteris and Tricholepidium are supported as monophyletic, the genus Paragramma is resurrected and the genera Lepisorus, Neolepisorus, Lemmaphyllum and Lepidomicrosorium are re-circumscribed. We proposed 14 new combinations, among which Caobangia is treated as a synonym of Lemmaphyllum. A key for identifying the recognized genera is presented. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 28–38.
The Osmundales (Royal Fern order) originated in the late Paleozoic and is the most ancient surviving lineage of leptosporangiate ferns. In contrast to its low diversity today (less than 20 species in six genera), it has the richest fossil record of any extant group of ferns. The structurally preserved trunks and rhizomes alone are referable to more than 100 fossil species that are classified in up to 20 genera, four subfamilies, and two families. This diverse fossil record constitutes an exceptional source of information on the evolutionary history of the group from the Permian to the present. However, inconsistent terminology, varying formats of description, and the general lack of a uniform taxonomic concept renders this wealth of information poorly accessible. To this end, we provide a comprehensive review of the diversity of structural features of osmundalean axes under a standardized, descriptive terminology. A novel morphological character matrix with 45 anatomical characters scored for 15 extant species and for 114 fossil operational units (species or specimens) is analysed using networks in order to establish systematic relationships among fossil and extant Osmundales rooted in axis anatomy. The results lead us to propose an evolutionary classification for fossil Osmundales and a revised, standardized taxonomy for all taxa down to the rank of (sub)genus. We introduce several nomenclatural novelties: (1) a new subfamily Itopsidemoideae (Guaireaceae) is established to contain Itopsidema, Donwelliacaulis, and Tiania; (2) the thamnopteroid genera Zalesskya, Iegosigopteris, and Petcheropteris are all considered synonymous with Thamnopteris; (3) 12 species of Millerocaulis and Ashicaulis are assigned to modern genera (tribe Osmundeae); (4) the hitherto enigmatic Aurealcaulis is identified as an extinct subgenus of Plenasium; and (5) the poorly known Osmundites tuhajkulensis is assigned to Millerocaulis. In addition, we consider Millerocaulis stipabonettiorum a possible member of Palaeosmunda and Millerocaulis estipularis as probably constituting the earliest representative of the (Todea-)Leptopteris lineage (subtribe Todeinae) of modern Osmundoideae.
Enlarged and highly repetitive plastome of Lagarostrobos and plastid phylogenomics of Podocarpaceae
Molecular Phylogenetics and Evolution
Podocarpaceae is the largest family in cupressophytes (conifers II), but its plastid genomes (plastomes) are poorly studied, with plastome data currently existing for only four of the 19 Podocarpaceous genera. In this study, we sequenced and assembled the complete plastomes from representatives of eight additional genera, including Afrocarpus, Dacrydium, Lagarostrobos, Lepidothamnus, Pherosphaera, Phyllocladus, Prumnopitys, and Saxegothaea. We found that Lagarostrobos, a monotypic genus native to Tasmania, has the largest plastome (151,496 bp) among any cupressophytes studied to date. Plastome enlargement in Lagarostrobos coincides with increased intergenic spacers, repeats, and duplicated genes. Among the Podocarpaceae, Lagarostrobos has the most rearranged plastome, but its substitution rates are modest. Plastid phylogenomic analyses based on 81 plastid genes clarify the positions of previously conflicting Podocarpaceous genera. Tree topologies firmly support the division of Podocarpaceae into two sister clades: (1) the Prumnopityoid clade and (2) the clade containing Podocarpoid, Dacrydioid, Pherosphaera, and Saxegothaea. The Phyllocladus is nested within the Podocarpaceae, thus familial status of the monotypic Phyllocladaceae is not supported.
American Journal of Botany, 2003
The systematic position and generic differentiation of the morphologically and geographically outstanding tribe Epithemateae (Gesneriaceae) was analyzed using the rbcL/atpB-spacer and trnL-F intron-spacer regions of chloroplast DNA. In our analysis Epithemateae forms a strongly supported monophyletic clade (bootstrap [BS] ϭ 100%; jackknife [JK] ϭ 100%; decay index [DI] ϭ 12) and appears as sister to the rest of the paleotropical Gesneriaceae (ϭ subfamily Cyrtandroideae). The paleotropical Gesneriaceae form a monophyletic group (BS ϭ 88%; JK ϭ 85%; DI ϭ 3) that is sister to the neotropical Gesneriaceae (subfamily Gesnerioideae) plus Austral Gesneriaceae (subfamily Coronantheroideae) (BS ϭ 99%; JK ϭ 98%; DI ϭ 10). Within Epithemateae Rhynchoglossum is sister to the remaining Epithemateae (BS ϭ 97%; JK ϭ 96%; DI ϭ 12), in which Epithema is sister to a clade of two genera: Loxonia/ Stauranthera (BS ϭ 68%; JK ϭ 64%; DI ϭ 1), which form, together with Epithema, a sister clade (BS ϭ 85%; JK ϭ 83%; DI ϭ 2) to Whytockia and Monophyllaea. While the support for Loxonia and Stauranthera is moderate, the relationship of Whytockia and Monophyllaea is very strongly supported (BS ϭ 100%; JK ϭ 100%; DI ϭ 13). Apart from the somewhat surprising (but wellsubstantiated) isolated position of Rhynchoglossum, the results are in perfect accordance with the relationships worked out earlier on grounds of architectural and floral characters. Especially remarkable is the predicted coherence between the morphologically and geographically different genera Whytockia and Monophyllaea.