Search

We investigated selected species of the six subgenera of the genus Gunnera (Gunneraceae, Gunnerales, basal core eudicot) in order to confirm and to particularize for this taxon trait specifics on the occurrence of endodermis with Casparian bands in the organs of these representative species and to relate this endodermis to the structure of the vascular tissues. The general presence of endodermis with Casparian bands in stems and the vascular tissue arrangement varying from monostele to complex siphonostele is particularly specific for the genus. Petiolar endodermis was also characteristic and some species even had an endodermis surrounding veins in leaf lamina. Similarity of vascular tissue in stems and petioles in Gunnera species to seedless vascular plants is an interesting structural trait regarding phylogenic position of the genus. Only two species had simple, tetrarch to pentarch root vascular patterns common for eudicots, whereas all others were hexarch to multiple polyarch.

Comparative genome biology has unveiled the polyploid origin of all angiosperms and the role of recurrent polyploidization in the amplification of gene families and the structuring of genomes. Which species share certain ancient polyploidy events, and which do not, is ill defined because of the limited number of sequenced genomes and transcriptomes and their uneven phylogenetic distribution. Previously, it has been suggested that most, but probably not all, of the eudicots have shared an ancient hexaploidy event, referred to as the gamma triplication. In this study, detailed phylogenies of subfamilies of MADS-box genes suggest that the gamma triplication has occurred before the divergence of Gunnerales but after the divergence of Buxales and Trochodendrales. Large-scale phylogenetic and K(S)-based approaches on the inflorescence transcriptomes of Gunnera manicata (Gunnerales) and Pachysandra terminalis (Buxales) provide further support for this placement, enabling us to position the gamma triplication in the stem lineage of the core eudicots. This triplication likely initiated the functional diversification of key regulators of reproductive development in the core eudicots, comprising 75% of flowering plants. Although it is possible that the gamma event triggered early core eudicot diversification, our dating estimates suggest that the event occurred early in the stem lineage, well before the rapid speciation of the earliest core eudicot lineages. The evolutionary significance of this paleopolyploidy event may thus rather lie in establishing a species lineage that was resilient to extinction, but with the genomic potential for later diversification. We consider that the traits generated from this potential characterize extant core eudicots both chemically and morphologically.

Recent phylogenetic analyses revealed a grade with Ranunculales, Sabiales, Proteales, Trochodendrales, and Buxales as first branching eudicots, with the respective positions of Proteales and Sabiales still lacking statistical confidence. As previous analyses of conserved plastid genes remain inconclusive, we aimed to use and evaluate a representative set of plastid introns (group I: trnL; group II: petD, rpl16, trnK) and intergenic spacers (trnL-F, petB-petD, atpB-rbcL, rps3-rpl16) in comparison to the rapidly evolving matK and slowly evolving atpB and rbcL genes. Overall patterns of microstructural mutations converged across genomic regions, underscoring the existence of a general mutational pattern throughout the plastid genome. Phylogenetic signal differed strongly between functionally and structurally different genomic regions and was highest in matK, followed by spacers, then group II and group I introns. The more conserved atpB and rbcL coding regions showed distinctly lower phylogenetic information content. Parsimony, maximum likelihood, and Bayesian phylogenetic analyses based on the combined dataset of non-coding and rapidly evolving regions (>14 000 aligned characters) converged to a backbone topology of eudicots with Ranunculales branching first, a Proteales–Sabiales clade second, followed by Trochodendrales and Buxales. Gunnerales generally appeared as sister to all remaining core eudicots with maximum support. Our results show that a small number of intron and spacer sequences allow similar insights into phylogenetic relationships of eudicots compared to datasets of many combined genes. The non-coding proportion of the plastid genome thus can be considered an important information source for plastid phylogenomics.

A study of inflorescence and flower development in 12 species from four of the six subgenera of Gunnera (Gunneraceae) was carried out. In the species of subgenus Panke, initiation of floral apices along the partial inflorescences is acropetal but ends up in the late formation of a terminal flower, forming a cyme at maturity. The terminal flower is the largest and the most complete in terms of merosity and number of whorls and thus it is the most diagnostic in terms of species-level taxonomy. The lateral flowers undergo a basipetal gradient of organ reduction along the inflorescence, ranging from bisexual flowers (towards the distal region) to functionally (i.e. with staminodia) and structurally female flowers (towards the proximal region). Our results show that the terminal structure in Gunnera is a flower rather than a pseudanthium. The terminal flower is disymmetric, dimerous and bisexual, representing the common bauplan for Gunnera flowers. It has a differentiated perianth with two sepals and two alternate petals, the latter opposite the stamens and carpels. Comparisons with other members of the core eudicots with labile floral construction are addressed. We propose vegetative and floral putative synapomorphies for the sister-group relationship between Gunneraceae and Myrothamnaceae. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160, 262–283.

Recent molecular phylogenetic studies have shown that the Gunnerales, including Gunnera and Myrothamnus, form a clade, sister to other core eudicots. This systematic position puts Gunnera in a pivotal position regarding the evolution of morphological characters in eudicots. In particular, the dimerous flower organization of Gunnera has recently been considered precursor to the pentamerous condition found in most core eudicots. In order to understand the role of Gunnera in the flower evolution of core eudicots, new detailed studies of the flowers of Gunnera are needed. In this study, the flower ontogeny and morphology of Gunnera monoica and G. dentata as well as of G. macrophylla were studied with SEM. The new observations are compared with previous studies of G. manicata and G. herteri. The first four species have dimerous perfect or staminate flowers with two small median sepals, followed by two large petals opposite the two stamens. In G. herteri, the staminate flowers are reduced to single stamens. In ...

Phylogenetic relationships among many lineages of angiosperms have been clarified via the analysis of large molecular data sets. However, with a data set of three genes (18S rDNA, rbcL, and atpB), relationships among lineages of core eudicots (Berberidopsidales, Caryophyllales, Gunnerales, Santalales, Saxifragales, asterids, rosids) remain essentially unresolved. We added 26S rDNA sequences to a three-gene matrix for 201 eudicots (8430 base pair aligned nucleotides per taxon). Parsimony analyses provided moderate (84%) jackknife support for Gunnerales, which comprise the two enigmatic families Gunneraceae and Myrothamnaceae, as sister to all other core eudicots. This position of Gunnerales has important implications for floral evolution. A dimerous or trimerous perianth is frequently encountered in early-diverging eudicots (e.g., Buxaceae, Proteales, Ranunculales, Trochodendraceae), whereas in core eudicots, pentamery predominates. Significantly, dimery is found in Gunneraceae and perhaps Myrothamnaceae (the merosity of the latter has also been interpreted as labile). Parsimony reconstructions of perianth merosity demonstrate lability among early-diverging eudicots and further indicate that a dimerous perianth could be the immediate precursor to the pentamerous condition characteristic of core eudicots. Thus, the developmental canalization that yielded the pentamerous condition of core eudicots occurred after the node leading to Gunnerales.

PREMISE OF THE STUDY Recent analyses employing up to five genes have provided numerous insights into angiosperm phylogeny, but many relationships have remained unresolved or poorly supported. In the hope of improving our understanding of angiosperm phylogeny, we expanded sampling of taxa and genes beyond previous analyses. METHODS We conducted two primary analyses based on 640 species representing 330 families. The first included 25260 aligned base pairs (bp) from 17 genes (representing all three plant genomes, i.e., nucleus, plastid, and mitochondrion). The second included 19846 aligned bp from 13 genes (representing only the nucleus and plastid). KEY RESULTS Many important questions of deep-level relationships in the nonmonocot angiosperms have now been resolved with strong support. Amborellaceae, Nymphaeales, and Austrobaileyales are successive sisters to the remaining angiosperms (Mesangiospermae), which are resolved into Chloranthales + Magnoliidae as sister to Monocotyledoneae + [Ceratophyllaceae + Eudicotyledoneae]. Eudicotyledoneae contains a basal grade subtending Gunneridae. Within Gunneridae, Gunnerales are sister to the remainder (Pentapetalae), which comprises (1) Superrosidae, consisting of Rosidae (including Vitaceae) and Saxifragales; and (2) Superasteridae, comprising Berberidopsidales, Santalales, Caryophyllales, Asteridae, and, based on this study, Dilleniaceae (although other recent analyses disagree with this placement). Within the major subclades of Pentapetalae, most deep-level relationships are resolved with strong support. CONCLUSIONS Our analyses confirm that with large amounts of sequence data, most deep-level relationships within the angiosperms can be resolved. We anticipate that this well-resolved angiosperm tree will be of broad utility for many areas of biology, including physiology, ecology, paleobiology, and genomics.

Although Pentapetalae (comprising all core eudicots except Gunnerales) include ≈70% of all angiosperms, the origin of and relationships among the major lineages of this clade have remained largely unresolved. Phylogenetic analyses of 83 protein-coding and rRNA genes from the plastid genome for 86 species of seed plants, including new sequences from 25 eudicots, indicate that soon after its origin, Pentapetalae diverged into three clades: ( i ) a “superrosid” clade consisting of Rosidae, Vitaceae, and Saxifragales; ( ii ) a “superasterid” clade consisting of Berberidopsidales, Santalales, Caryophyllales, and Asteridae; and ( iii ) Dilleniaceae. Maximum-likelihood analyses support the position of Dilleniaceae as sister to superrosids, but topology tests did not reject alternative positions of Dilleniaceae as sister to Asteridae or all remaining Pentapetalae. Molecular dating analyses suggest that the major lineages within both superrosids and superasterids arose in as little as 5 million years. This phylogenetic hypothesis provides a crucial historical framework for future studies aimed at elucidating the underlying causes of the morphological and species diversity in Pentapetalae.

A close relationship between Berberidopsidaceae and Aextoxicaceae has never been considered until gene sequence studies provided strong support for a relationship between them (see family treatments). In the four-gene analysis of eudicots (Soltis et al. 2003), Gunnerales and subsequently Berberidopsidales are sister to all other core eudicots, the latter being strongly supported by molecular data and isolated from all other clades (Fig. 1). Aextoxicum has long been known for its peculiar wood anatomy, particularly the high number of bars of the vessel element perforations. A recent study by Carlquist (2003) has revealed many important similarities in the wood anatomy of the two families, although these are plesiomorphic. Pollen grains are relatively small and tricolpate to indistinctly colporate. The two families share encyclocytic stomata (Soltis et al. 2005), a rare character in angiosperms, stout filaments, and a ring of vascular bundles in the petiole (Judd and Olmstead 2004).