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Phylogenetic relationships and evolution of extant Horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F)

Equisetumis a small and morphologically distinct genus with a rich fossil record. Two subgenera have been recognized based principally on stomatal position and stem branching: subg.Equisetum(eight species; superficial stomates; stems branched) and subg.Hippochaete(seven species; sunken stomates; ste...

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Published in:International journal of plant sciences 2003-09, Vol.164 (5), p.737-751
Main Authors: Des Marais, D.L, Smith, A.R, Britton, D.M, Pryer, K.M
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Pryer, K.M
description Equisetumis a small and morphologically distinct genus with a rich fossil record. Two subgenera have been recognized based principally on stomatal position and stem branching: subg.Equisetum(eight species; superficial stomates; stems branched) and subg.Hippochaete(seven species; sunken stomates; stems generally unbranched). Prior attempts at understandingEquisetumsystematics, phylogeny, and character evolution have been hampered by the high degree of morphological plasticity in the genus as well as by frequent hybridization among members within each subgenus. We present the first explicit phylogenetic study ofEquisetum, including all 15 species and two samples of one widespread hybrid,Equisetum×ferrissii, based on a combined analysis of two chloroplast markers,rbcLandtrnL‐F. Our robustly supported phylogeny identifies two monophyletic clades corresponding to the two subgenera recognized by earlier workers. The phylogenetic placement ofEquisetum bogotense, however, is ambiguous. In maximum likelihood analyses, it allies with subg.Hippochaeteas the most basal member, while maximum parsimony places it as sister to the rest of the genus. A consensus phylogeny from the two analyses is presented as a basal trichotomy (E. bogotense, subg.Hippochaete, subg.Equisetum), and morphological character evolution is discussed. We detected rate heterogeneity in therbcLlocus between the two subgenera that can be attributed to an increased rate of nucleotide substitution (transversions) in subg.Hippochaete. We calculated molecular‐based age estimates using the penalized likelihood approach, which accounts for rate heterogeneity and does not assume a molecular clock. TheEquisetumcrown group appears to have diversified in the early Cenozoic, whereas the Equisetaceae total group is estimated to have a Paleozoic origin. These molecular‐based age estimates are in remarkable agreement with current interpretations of the fossil record.
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Two subgenera have been recognized based principally on stomatal position and stem branching: subg.Equisetum(eight species; superficial stomates; stems branched) and subg.Hippochaete(seven species; sunken stomates; stems generally unbranched). Prior attempts at understandingEquisetumsystematics, phylogeny, and character evolution have been hampered by the high degree of morphological plasticity in the genus as well as by frequent hybridization among members within each subgenus. We present the first explicit phylogenetic study ofEquisetum, including all 15 species and two samples of one widespread hybrid,Equisetum×ferrissii, based on a combined analysis of two chloroplast markers,rbcLandtrnL‐F. Our robustly supported phylogeny identifies two monophyletic clades corresponding to the two subgenera recognized by earlier workers. The phylogenetic placement ofEquisetum bogotense, however, is ambiguous. In maximum likelihood analyses, it allies with subg.Hippochaeteas the most basal member, while maximum parsimony places it as sister to the rest of the genus. A consensus phylogeny from the two analyses is presented as a basal trichotomy (E. bogotense, subg.Hippochaete, subg.Equisetum), and morphological character evolution is discussed. We detected rate heterogeneity in therbcLlocus between the two subgenera that can be attributed to an increased rate of nucleotide substitution (transversions) in subg.Hippochaete. We calculated molecular‐based age estimates using the penalized likelihood approach, which accounts for rate heterogeneity and does not assume a molecular clock. TheEquisetumcrown group appears to have diversified in the early Cenozoic, whereas the Equisetaceae total group is estimated to have a Paleozoic origin. 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subjects Biological taxonomies
biomarkers
chloroplasts
Equisetum
Equisetum bogotense
Evolution
Flowers & plants
Fossils
Gametophytes
Genetics
Hippochaete
Horsetails
Hybridity
molecular sequence data
molecular systematics
nucleotide sequences
Parsimony
Phylogeny
plant proteins
rbcL gene
rbcL protein
ribulose-bisphosphate carboxylase
sequence alignment
sequence analysis
Stomata
Taxa
trnF gene
trnL gene
trnL protein
We they distinction
title Phylogenetic relationships and evolution of extant Horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F)
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