Phylogenetic relationships among the four major lineages of land plants (liverworts, mosses, hornworts, and vascular plants) remain vigorously contested; their resolution is essential to our understanding of the origin and early evolution of land plants. We analyzed three different complementary data sets: a multigene supermatrix, a genomic structural character matrix, and a chloroplast genome sequence matrix, using maximum likelihood, maximum parsimony, and compatibility methods. Analyses of all three data sets strongly supported liverworts as the sister to all other land plants, and analyses of the multigene and chloroplast genome matrices provided moderate to strong support for hornworts as the sister to vascular plants. These results highlight the important roles of liverworts and hornworts in two major events of plant evolution: the water-to-land transition and the change from a haploid gametophyte generation-dominant life cycle in bryophytes to a diploid sporophyte generation-dominant life cycle in vascular plants. This study also demonstrates the importance of using a multifaceted approach to resolve difficult nodes in the tree of life. In particular, it is shown here that densely sampled taxon trees built with multiple genes provide an indispensable test of taxon-sparse trees inferred from genome sequences.alternation of generations ͉ hornworts ͉ liverworts ͉ phylogeny ͉ taxon sampling T he origin and early evolution of land plants (embryophytes) during the mid-Ordovician to lower Silurian (480-430 million years ago) initiated the establishment of the modern terrestrial ecosystems and fundamentally altered the course of evolution of life on earth. Two important events marked this period of unprecedented innovation in plant evolution: the massive colonization of the land by plants descended from charophyte algae and the change of the dominant generation in the plant life cycle from a haploid gametophyte to a diploid sporophyte (1-5). The first event opened a vastly underexplored niche of high-intensity solar radiation and abundant CO 2 to photosynthetic life. The second event conferred on plants two abilities to adapt to a life in a water-deficient and UV-abundant terrestrial environment. One is the ability to produce a large number of genetically diverse gametes to ensure fertilization on land where sperm locomotion is hindered, and the other is the ability to mask deleterious mutations through the dominantrecessive interaction of alleles, thus allowing a large number of alleles to persist in the gene pool (2-4). Our understanding of these events hinges on our knowledge of relationships between the organisms involved in these major evolutionary transitions. Despite numerous studies using diverse approaches analyzing morphological and͞or molecular characters, relationships among early land plants remain controversial (5-19). Fossil evidence, although increasingly improved, has not helped to resolve the issues decisively (20,21).A multitude of phenomena characterizing diversification of many major clades o...
Ford, C. S., Ayres, K. L., Toomey, N., Haider, N., Stahl, J. V., Kelly, L. J., Wikstrom, N., Hollingsworth, P. M., Duff, R. J., Hoot, S. B., Cowan, R. S., Chase, M. W., Wilkinson, M. J. (2009). Selection of candidate coding DNA barcoding regions for use on land plants. Botanical Journal of the Linnean Society, 159, (1), 1-11. Sponsorship: Alfred P. Sloan Foundation Gordon and Betty Moore Foundation IMPF: 00.98 RONO: 00An in silico screen of 41 of the 81 coding regions of the Nicotiana plastid genome generated a shortlist of 12 candidates as DNA barcoding loci for land plants. These loci were evaluated for amplification and sequence variation against a reference set of 98 land plant taxa. The deployment of multiple primers and a modified multiplexed tandem polymerase chain reaction yielded 85?94% amplification across taxa, and mean sequence differences between sister taxa of 6.1 from 156 bases of accD to 22 from 493 bases of matK. We conclude that loci should be combined for effective diagnosis, and recommend further investigation of the following six loci: matK, rpoB, rpoC1, ndhJ, ycf5 and accD.Peer reviewe
A widely held view of land plant relationships places liverworts as the first branch of the land plant tree, whereas some molecular analyses and a cladistic study of morphological characters indicate that hornworts are the earliest land plants. To help resolve this conflict, we used parsimony and likelihood methods to analyze a 6, 095-character data set composed of four genes (chloroplast rbcL and small-subunit rDNA from all three plant genomes) from all major land plant lineages. In all analyses, significant support was obtained for the monophyly of vascular plants, lycophytes, ferns (including PSILOTUM: and EQUISETUM:), seed plants, and angiosperms. Relationships among the three bryophyte lineages were unresolved in parsimony analyses in which all positions were included and weighted equally. However, in parsimony and likelihood analyses in which rbcL third-codon-position transitions were either excluded or downweighted (due to apparent saturation), hornworts were placed as sister to all other land plants, with mosses and liverworts jointly forming the second deepest lineage. Decay analyses and Kishino-Hasegawa tests of the third-position-excluded data set showed significant support for the hornwort-basal topology over several alternative topologies, including the commonly cited liverwort-basal topology. Among the four genes used, mitochondrial small-subunit rDNA showed the lowest homoplasy and alone recovered essentially the same topology as the multigene tree. This molecular phylogeny presents new opportunities to assess paleontological evidence and morphological innovations that occurred during the early evolution of terrestrial plants.
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