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Increasingly robust understanding of angiosperm phylogeny allows more secure reconstruction of the flower in the most recent common ancestor of extant angiosperms and its early evolution. The surprising emergence of several extant and fossil taxa with simple flowers near the base of the angiosperms-Chloranthaceae, Ceratophyllum, Hydatellaceae, and the Early Cretaceous fossil Archaefructus (the last three are water plants)-has brought a new twist to this problem. We evaluate early floral evolution in angiosperms by parsimony optimization of morphological characters on phylogenetic trees derived from morphological and molecular data. Our analyses imply that Ceratophyllum may be related to Chloranthaceae, and Archaefructus to either Hydatellaceae or Ceratophyllum. Inferred ancestral features include more than two whorls (or series) of tepals and stamens, stamens with protruding adaxial or lateral pollen sacs, several free, ascidiate carpels closed by secretion, extended stigma, extragynoecial compitum, and one or several ventral pendent ovule(s). The ancestral state in other characters is equivocal: e.g., bisexual vs. unisexual flowers, whorled vs. spiral floral phyllotaxis, presence vs. absence of tepal differentiation, anatropous vs. orthotropous ovules. Our results indicate that the simple flowers of the newly recognized basal groups are reduced rather than primitively simple.

The angiosperm order Malpighiales includes ∼16,000 species and constitutes up to 40% of the understory tree diversity in tropical rain forests. Despite remarkable progress in angiosperm systematics during the last 20 y, relationships within Malpighiales remain poorly resolved, possibly owing to its rapid rise during the mid-Cretaceous. Using phylogenomic approaches, including analyses of 82 plastid genes from 58 species, we identified 12 additional clades in Malpighiales and substantially increased resolution along the backbone. This greatly improved phylogeny revealed a dynamic history of shifts in net diversification rates across Malpighiales, with bursts of diversification noted in the Barbados cherries (Malpighiaceae), cocas (Erythroxylaceae), and passion flowers (Passifloraceae). We found that commonly used a priori approaches for partitioning concatenated data in maximum likelihood analyses, by gene or by codon position, performed poorly relative to the use of partitions identified a posteriori using a Bayesian mixture model. We also found better branch support in trees inferred from a taxon-rich, data-sparse matrix, which deeply sampled only the phylogenetically critical placeholders, than in trees inferred from a taxon-sparse matrix with little missing data. Although this matrix has more missing data, our a posteriori partitioning strategy reduced the possibility of producing multiple distinct but equally optimal topologies and increased phylogenetic decisiveness, compared with the strategy of partitioning by gene. These approaches are likely to help improve phylogenetic resolution in other poorly resolved major clades of angiosperms and to be more broadly useful in studies across the Tree of Life. M alpighiales are one of the most surprising clades discovered in broad molecular phylogenetic studies of the flowering plants (1-3). The order contains ∼16,000 species and 42 families (2, 3) that exhibit remarkable morphological and ecological diversity. A few examples include cactus-like succulents (Euphorbiaceae), epiphytes (Clusiaceae), holoparasites (Rafflesiaceae), submerged aquatics (Podostemaceae), and windpollinated trees (temperate Salicaceae). The order is ecologically important: species in Malpighiales constitute up to 40% of the understory tree diversity in tropical rain forests worldwide (4). They also include many economically important species, such as Barbados nut (Jatropha curcas L., Euphorbiaceae), cassava (Manihot esculenta Crantz, Euphorbiaceae), castor bean (Ricinus communis L., Euphorbiaceae), coca (Erythroxylum coca Lam., Erythroxylaceae), flax (Linum usitatissimum L., Linaceae), the poplars (Populus spp., Salicaceae), and the rubber tree (Hevea brasiliensis Müll. Arg., Euphorbiaceae). Partially for this reason, genomic resources for Malpighiales are growing at a rapid pace and include whole-genome sequencing projects completed or near completion for Barbados nut (5), cassava, castor bean (6), flax, and poplar (7). Thus, a resolved phylogeny of Malpighiales is critical not only for evol...

Recent advances in molecular phylogenetics and a series of important palaeobotanical discoveries have revolutionized our understanding of angiosperm diversification. Yet, the origin and early evolution of their most characteristic feature, the flower, remains poorly understood. In particular, the structure of the ancestral flower of all living angiosperms is still uncertain. Here we report model-based reconstructions for ancestral flowers at the deepest nodes in the phylogeny of angiosperms, using the largest data set of floral traits ever assembled. We reconstruct the ancestral angiosperm flower as bisexual and radially symmetric, with more than two whorls of three separate perianth organs each (undifferentiated tepals), more than two whorls of three separate stamens each, and more than five spirally arranged separate carpels. Although uncertainty remains for some of the characters, our reconstruction allows us to propose a new plausible scenario for the early diversification of flowers, leading to new testable hypotheses for future research on angiosperms.

This article traces research on floral symmetry back to its beginnings. It brings together recent advances from different fields that converge in floral symmetry and new unpublished material on diversity and development of floral symmetry. During floral development, symmetry may change: monosymmetric flowers may have a polysymmetric early phase; polysymmetric flowers may have a monosymmetric or even asymmetric early phase; more than one symmetry change is also possible. In Lamiales s.l. (comprising the model plant Antirrhinum, where the cycloidea gene produces monosymmetric flowers with the adaxial side of the androecium reduced), taxa also occur in which the androecium is reduced on both sides, adaxial and abaxial. As a trend in asymmetric flowers, enantiomorphy (with two mirror-image morphs) at the level of individuals seems to occur only in groups in which the flowers are predominantly of a relatively simple construction. In contrast, one morph is fixed at the level of species or higher taxa in groups with more complicated flowers. This is indicated by the apparent lack of enantiomorphy in corolla contortion in asterids but its predominance in rosids with contort flowers, or by the apparent lack of enantiomorphy in the pollination organs of asymmetric flowers in Faboideae but its presence in asymmetric flowers in Caesalpinioideae. To study the evolution of the diverse symmetry patterns, a concerted approach from different fields including molecular developmental genetics, pollination biology, and comparative diversity research is necessary.

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Angiosperms and their flowers have greatly diversified into an overwhelming array of forms in the past 135 million years. Diversification was shaped by changes in climate and the biological environment (vegetation, interaction with other organisms) and by internal structural constraints and potentials. This review focuses on the development and structural diversity of flowers and structural constraints. It traces floral diversification in the different organs and organ complexes (perianth, androecium, gynoecium) through the major clades of extant angiosperms. The continuously improved results of molecular phylogenetics provide the framework for this endeavor, which is necessary for the understanding of the biology of the angiosperms and their flowers. Diversification appears to work with innovations and modifications of form. Many structural innovations originated in several clades and in special cases could become key innovations, which likely were hot spots of diversification. Synorganization between organs was an important process to reach new structural levels, from which new diversifications originated. Complexity of synorganization reached peaks in Orchidaceae and Apocynaceae with the independent evolution of pollinaria. Such a review throughout the major clades of angiosperms also shows how superficial and fragmentary our knowledge on floral structure in many clades is. Fresh studies and a multidisciplinary approach are needed.