Archaefructus

Abstract: 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.

Abstract: The early history of flowering plants (angiosperms) is contentious, but a steady stream of fossil discoveries from the Early Cretaceous of China is beginning to fill in some of the gaps in our knowledge. The latest of these, found in the Lower Cretaceous Yixian Formation, is a fossil that dates to between 124.4 million and 122.9 million years old. It is particularly informative because this angiosperm is a eudicot — the relatively derived group that dominates the flowering plants in the world today — and is also possibly a member of the Ranunculaceae. The fossil confirms the presence of the eudicots at this time and documents an early burst of angiosperm evolution. The early history of flowering plants (angiosperms) is contentious. However, many fossils attributable to flowering plants have been found in the Early Cretaceous of China. The plant reported in this study goes a step further: not only is it an angiosperm, but it is a member of a relatively derived group, the eudicots, and possibly a member of the Ranunculaceae, an extant family of plants. This indicates that angiosperm evolution probably got into its stride even earlier than the Early Cretaceous. The current molecular systematics of angiosperms1 recognizes the basal angiosperms and five major angiosperm lineages: the Chloranthaceae, the magnoliids, the monocots, Ceratophyllum and the eudicots, which consist of the basal eudicots and the core eudicots2. The eudicots form the majority of the angiosperms in the world today. The flowering plants are of exceptional evolutionary interest because of their diversity of over 250,000 species and their abundance as the dominant vegetation in most terrestrial ecosystems, but little is known of their very early history. In this report we document an early presence of eudicots during the Early Cretaceous Period. Diagnostic characters of the eudicot fossil Leefructus gen. nov. include simple and deeply trilobate leaves clustered at the nodes in threes or fours, basal palinactinodromous primary venation, pinnate secondary venation, and a long axillary reproductive axis terminating in a flattened receptacle bearing five long, narrow pseudo-syncarpous carpels. These morphological characters suggest that its affinities are with the Ranunculaceae, a basal eudicot family. The fossil co-occurs with Archaefructus sinensis3 and Hyrcantha decussata4 whereas Archaefructus liaoningensis5 comes from more ancient sediments. Multiple radiometric dates of the Lower Cretaceous Yixian Formation place the bed yielding this fossil at 122.6–125.8 million years old6,7,8. The earliest fossil records of eudicots are 127 to 125 million years old, on the basis of pollen9,10. Thus, Leefructus gen. nov. suggests that the basal eudicots were already present and diverse by the latest Barremian and earliest Aptian.

Abstract: Recently, whether Archaefructus has bisexual multi-parted flowers or just inflorescences of unisexual flowers, and whether it is ancestral to all angiosperms or a derived eudicot have been debated. Here, from the same Yixian Formation, NE China, we report a new Archaefructus species, A. eoflora sp. nov., with the generic characteristics of dissected leaves and bisexual reproductive axes. It is entirely preserved with roots, rhizome, shoots and protogynous reproductive organs at different developmental stages. Its lateral and main fertile shoots form a pseudo-indeterminate pattern, while the reproductive branches on the main shoot form a cymose inflorescence. Subtended by 1-2 bract-like leaves, the section of stamen clusters changes into a much shorter section of carpels that have one row of orthotropous ovules. Significantly, one cluster bearing two carpels and one stamen between the two sections demonstrates a true bisexual flower, an important step of the origin of floral bisexuality, and the homology between the stamen and carpel. The complex reproductive axes represent a mix between flowers and inflorescences, and suggest that A. eoflora sp. nov. possesses the potential to evolve into a variety of diverse flower types as found in modern basal and early fossil angiosperms. A comparison with other two species also leads to a revision of the generic diagnosis.

Abstract: The origin of angiosperms has been under debate since the time of Darwin. While there has been much speculation in past decades about pre-Cretaceous angiosperms, including Archaefructus, these reports are controversial. The earliest reliable fossil record of angiosperms remains restricted to the Cretaceous, even though recent molecular phylogenetic studies suggest an origin for angiosperms much earlier than the current fossil record. In this paper, after careful SEM and light microscopic work, we report fossils with angiospermous traits of the Jurassic age. The fossils were collected from the Haifanggou Formation (middle Jurassic) in western Liaoning, northeast China. They include two female structures and an associated leaf on the same slab. One of the female structures is physically connected to the apex of a short shoot. The female organs are borne in pairs on short peduncles that are arranged along the axis of the female structure. Each of the female organs has a central unit that is surrounded by an envelope with characteristic longitudinal ribs. Each central unit has two locules completely separated by a vertical septum. The apex of the central unit is completely closed. The general morphology places these fossils into the scope of Schmeissneria, an early Jurassic genus that was previously attributed to Ginkgoales. Because the closed carpel is a character only found in angiosperms, the closed apex of the central unit suggests the presence of angiospermy in Schmeissneria. This angiospermous trait implies either a Jurassic angiosperm or a new seed plant group parallel to angiosperms and other known seed plants. As an angiosperm, the Liassic age (earliest Jurassic) of Schmeissneria microstachys would suggest an origin of angiosperms during the Triassic. Although still uncertain, this could have a great impact on our perspective of the history, diversity and systematics of seed plants and angiosperms.