The mitochondrial phylogeny of land plants shows support for Setaphyta under composition-heterogeneous substitution models

Congruence among analyses of plant genomic data partitions (nuclear, chloroplast and mitochondrial) is a strong indicator of accuracy in plant molecular phylogenetics. Recent analyses of both nuclear and chloroplast genome data of land plants (embryophytes) have, controversially, been shown to support monophyly of both bryophytes (mosses, liverworts, and hornworts) and tracheophytes (lycopods, ferns, and seed plants), with mosses and liverworts forming the clade Setaphyta. However, relationships inferred from mitochondria are incongruent with these results, and typically indicate paraphyly of bryophytes with liverworts alone resolved as the earliest-branching land plant group. Here, we reconstruct the mitochondrial land plant phylogeny from a newly compiled data set. When among-lineage composition heterogeneity is accounted for in analyses of codon-degenerate nucleotide and amino acid data, the clade Setaphyta is recovered with high support, and hornworts are supported as the earliest-branching lineage of land plants. These new mitochondrial analyses demonstrate partial congruence with current hypotheses based on nuclear and chloroplast genome data, and provide further incentive for revision of how plants arose on land.

Response to Comment on “A promiscuous intermediate underlies the evolution of LEAFY DNA binding specificity”

Brunkard et al. propose that the identification of novel LEAFY sequences contradicts our model of evolution through promiscuous intermediates. Based on the debate surrounding land plant phylogeny and on our analysis of these interesting novel sequences, we explain why there is no solid evidence to disprove our model.

Ultrastructure of the spermatozoid of Lycopodiella lateralis (Lycopodiaceae)

The spermatozoid of Lycopodiella lateralis (R. Br.) B. Øllgaard is ovoid and biflagellated and contains little cytoplasm. A large, rounded, condensed nucleus occupies the central region of the cell. At the rear of the cell is a cluster of starch-containing plastids associated with numerous elongated mitochondria and small pockets of remnant cytoplasm. The multilayered structure coils laterally around the cell anterior for just over one revolution. An anterior mitochondrion underlies the multilayered structure over its entire length and several smaller mitochondria line the inner anterior coil. Some 150 spline microtubules extend from the multilayered structure and coil posteriorly at approximately a 45° angle to the longitudinal axis of the cell. Microtubules progressively increase in length from the margins of the multilayered structure to the center and thus only a central core of approximately 30 microtubules encircles the organelles at the base of the cell. The two basal bodies are parallel and staggered in their position over the multilayered structure and are separated by about 80° around the circumference of the cell anterior. The flagella emerge in parallel and coil for nearly two revolutions. Comparisons among Lycopodiella, Palhinhaea, and Lycopodium provide the basis for evolutionary inferences associated with modifications in spermatozoid structure such as changes in cell size, whole cell coiling, and distance between basal bodies. Phylogenetic analysis of male gametogenesis suggests that Lycopodiella is part of a monophyletic lycopsid assemblage near the base of the vascular plants. Within this clade, Lycopodiella is most closely related to Palhinhaea, with Lycopodium, Phylloglossum, and Selaginella forming a sister clade. Key words: Lycopodiella, Lycopodium, spermatozoid, land plant phylogeny, locomotory apparatus, ultrastructure.

Occurrence and phylogenetic significance of monoplastidic meiosis in liverworts

Monoplastidic meiosis is reported for the first time in three seemingly unrelated liverworts, namely Blasia pusilla (Metzgeriales), Monoclea gottschei (Monocleales), and Haplomitrium blumei (Haplomitriales). A second species of Haplomitrium, H. hookeri, is polyplastidic as previously reported. All three taxa represent isolated relicts of ancient liverwort lineages. Monoplastidy in these hepatics is evident in archesporial tissue and is maintained through successive sporogenous cell generations. In archesporial mitosis, the single plastid divides and the two resultant plastids are precisely positioned so that one is inherited by each daughter cell. In the nascent spore mother cell, the solitary plastid undergoes two successive divisions and the resulting four plastids become positioned in a tetrahedral arrangement. Concomitantly, the sporocyte assumes a quadrilobed shape, which is less exaggerated in Monoclea, and a single large plastid is situated in each lobe. Details of plastid ultrastructure and morphology vary slightly among the three taxa. Evidence is presented that Blasia and Monoclea share a common ancestry and represent pivotal taxa in the evolution of the two main lines of liverworts. Haplomitrium is suggested to occupy a more basal position in bryophyte phylogeny. Monoplastidy in meiosis of liverworts links the charophytes, the three bryophyte clades, and the lycopsid pteridophytes and supports a monophyletic interpretation of land plant phylogeny. Key words: chloroplast, liverwort, meiosis, monoplastidy, phylogeny, sporogenesis.