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Evolution of stem- and crown-group flowering plants is discussed from molecular-systematic-, floral tool kit-, and paleobotanical research perspectives in this third of three essays on the origin of angiosperms. This statement by Wing and Boucher (page 380, 1998) is probably incorrect: "Despite the singular ecological significance and species diversity of angiosperms, they are not in a genealogical sense one of the major branches of land plants and did not originate with other major land plant clades (e.g. Some of the high points on floral evo-devo of eudicots with bearing on the greater question of the timing of the origin of floral organs are published in recent works by Hileman and Irish (2009), Korotkova et al. Additional permineralized fossil material of Sanmiguelia is probably needed to better understand the anatomy of reproduction and whole plant morphology. Uncanny similarities of early Mesozoic seed plant Sanmiguelia lewisii (Cornet 1986, 1989) with Paleozoic Vojnovskyales pointed out by Crane (1985) require confirmation by phylogenetic analysis of reproductive and vegetative characters gleaned from detailed anatomical studies of more fossilized remains to be collected. The Archaemagnoliidae is lumped with the Magnoliidae. Molecular-phylogenetic studies suggest that differentiation of flowering plants into a stem and crown group of the Mesozoic Era is feasible (Hilu et al.

I also bring back to life monocotyledonous elements of a ghost lineage of flowering plants traced from Norian sanmiguelias. Paleoecologies of these ancient stem-group angiosperm populations were not "dark and disturbed," or "wet and wild," or explainable by any other nonsensical and sophomorical pairing of adjectives. lycopsids, ferns, conifers, cycads, ginkgos) during the middle or late Paleozoic." Absence of paleobotanical data is not a substitute for fact when dealing with a probable ghost lineage due to insufficient sampling, especially in view of several molecular-phylogenetic studies estimating divergences of the flowering plant crown more than 220 MYA, which were events centered in the middle of the Triassic Period (Stephen A. Ovuliferous inflorescences first described as Axelrodia burgeri, polleniferous inflorescences named Synangispadixis tidwellii, flowers with ovuliferous units and polleniferous units, megasporophylls as carpels, synangia as anthers, bracts, and bitegmic ovules were described and discussed by Cornet (1986, 1989). This is an unstudied chronocline with potentially profound implications toward the idea of paraphyletic transitions in diverging seed plants at the base of the angiosperm stem(s) that straddle the PT. Triassic angiosperm fossils of detached "dicot-like" leaves described as Pannaulika triassica are known (Cornet 1993). Fossils of several other enigmatic flowering plants have been recovered from Mesozoic rocks but reproductive details and the morphology of whole plants are unclear due to problems with poor preservation and uncertain stratigraphic control (Müller 1981, G. There is a significant increase in the number of orders and genera of fossil flowering plants by the Aptian Age of the Gallic Epoch of the Cretaceous Period, based on data in Table 5.

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(2009) are probably better working seed plant phylogenies than the alternative trees computed by Bateman et al. (4[b], Implications for the origin of eudicots, 2016). Basinger and Dilcher's "Rose Creek Flower" and Caliciflora mauldiniensis (3. 2016) figure prominently in discussions of rosid phylogenies and the evolution of floral pentamery. (A) Two orthogonal views of LFY-C helices α1 - α3 bound to their DNA target site [red] superimposed with the three helical bundle core of the N-terminal subdomain of the paired domain of Drosophila Prd [blue, PDB-id: 1pdn]. Calibrating the timing of this split together with the other Great Late Paleozoic Seed Plant Divergences, using guide fossils, will be important exercises in future combined molecular- and morphological-phylogenetic analyses. Employing refined methods to estimate rates of divergence of flowering plants, Charles Bell and colleagues (2010) publish a milestone paper based on more than 500 extant taxa and 35 calibration points. Based on tool kit studies of homeodomain proteins there might be indirect evidence that populations of seed plants with flowers existed prior to the PT. (2016) suggest another scenario, which is at odds with another point of view expressed by Becker (2016). Biochemical studies of cis-acting TFs (Becker 2016) and anatomical evidence of the paleobiology of auxin-mediated secondary growth (Rothwell and Lev-Yadun 2005, Rothwell et al. Peter Endress (2015) reviews the developmental morphology of the angiosperm carpel. the supposed source of shed Afropollis), Bomfleur and coauthors (op. Doyle (2008), former iterations of this seed plant phylogeny, and some hypotheses billed as "theories" (Frohlich 2002), are probably incorrect. "We have examined herein different methodological approaches (i.e. This rather common Triassic fossil of southwestern North America is remarkably similar to the Paleozoic Vojnovskyales (page 779, Crane 1985). Possibly, but this determination must be validated by way of paleobotanical evidence yet to be mined from the rocks. Absence of fossilized remains of flowering plants in the stratigraphic record is not data. Any classification based on a single organ has a greater potential for error than one based on a variety of organs ... Integers in Table 5 represent the total number of taxonomic orders and genera (in parentheses) for each of Cronquist's subclasses of flowering plants.

(2016), Rothwell and Stockey (2016) and Rothwell et al. The rock contains an indeterminate eudicot fossil flower, which is the same "Rose Creek Flower" discussed by Friis et al. Dilcher (1984), Ancient bisexual flowers, Science 224: 511-513. (2008), "Comparison of LFY-C with paired and homeodomain DNA binding. "Did insect pollination cause increased seed plant diversity? These questions, among others (Berendse and Scheffer 2009, Crepet and Niklas 2009) should now be discussed from a late Paleozoic temporal perspective based on evidence presented in the first and second essays. There is growing consensus among some molecular systematists and paleobotanists on the existence of a 160 million year old angiosperm ghost lineage rooted at the angiosperm-gymnosperm split roughly 300 MYA, prior to the end-Permian extinction. Vivian Irish (2006) provides a road map to diversification of the angiosperm clade from the perspective of evo-devo of floral homeotic genes, their phenotypic expression, and molecular phylogenies. Specifically, our results indicate that the heterodimerization between DEF-like and GLO-like proteins was already present in the [most common recent ancestor] MRCA of extant angiosperms and was virtually never rewired" (page 1438, Discussion, Conservation of the MADS-domain protein interaction pattern during angiosperm evolution, Melzer et al. Most of the explosive radiation of floral diversity in basal lineages of flowering plants is explained by duplication and diversification of the MIKC-type MADS-box family of genes (P. The reconstructed physiology and ecology of the Petriellales matches this life form to such detail that we suggest these unusual gymnosperms may represent convergent ecological analogues of early flowering plants" (page 1074, Discussion, Ecology and Paleoenvironment, Bomfleur et al. Taking into account the morphology of reproductive short shoots of Winteraceae (i.e. Hochuli, Palaeontological Institute and Museum, University of Zürich, Zürich, Switzerland. I completely reject the assertion by some workers that angiosperms first appeared in the Cretaceous Period (Feild and Arens 2007). "Angiosperm phylogeny is riddled with examples of convergent morphologies ...

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Specimen IU15713-3429 was first photographed by the author in 1981 with the permission of Professor David L. The same specimen was illustrated in Figure 1 [as "D"] on Page 512 of J. A Mesozoic radiation of angiosperms is cast within the framework of the Angiosperm Phylogeny Group proposed classification (APG III 2009, Chase and Reveal 2009, APG IV 2016). (B) Superposition with the homeodomain of Drosophila engrailed bound to DNA [yellow, PBD-id: 1hdd], where the centre of recognition helix α3 inserts into the major groove." Reprinted by permission from Macmillan Publishers Ltd: The European Molecular Biology Organization (EMBO) Journal, Hamès, C., D. A possible paraphyletic Paleozoic origin of angiosperms is incongruent with proposals by Leebens-Mack et al. Ancestors of putative paraphyletic grades of angiosperms might have been Permo-carboniferous or Permo-triassic gymnosperms with hermaphroditic (bisexual) strobili. 2008) suggests deep conservation of the developmental tool kit of vascular plants. "While the amazing conservation of the major floral identity [ABCDE] program is being unravelled by analysing floral homeotic gene function and expression, we are only just beginning to understand the evolution of the gene network governing the organ identity genes ... cit.) should understand that petriellalean cupules are incompatible with tool kit models of floral development from shoot apical meristems (SAMs). We anticipate that the evident question-whether beyond the mere ecological similarity there may be phylogenetic relationships linking Petriellales with angiosperms-will be answered once more detailed information about their reproductive biology becomes available" (page 1074, Discussion, Ecology and Paleoenvironment, Bomfleur et al. Caytoniales, Corystospermales, Doyleales, and Petriellales probably had nothing to do with the evolution of stem-group flowering plants or the origin of angiosperms. Is the bitegmic ovule an angiosperm-specific character? Conversely, can ategmic ovules develop by fusion of integuments? fossil-based, molecular, phylogenetic and paleobiogeographic studies) and current viewpoints about the explosive Cretaceous diversification of angiosperms. Of all the enigmatic seed plants of the early Mesozoic Era, Sanmiguelia lewisii has attracted the most attention by students of angiosperm paleobiology (S. Crane (1985) suggested that some populations of late Paleozoic Vojnovskyales might have survived the end-Permian extinction reappearing in the Triassic rock record as the seed plant Sanmiguelia. The problem is also semantic as expressed in conflicting opinions on the definition of angiosperms, flowers, and from opposing ideas on supposed character homologies with organs of Paleozoic seed plants. Table 5 summarizes the stratigraphic distribution and microfossil, megafossil, and mesofossil history of the subclasses of flowering plants according to Cronquist (1981). Although one solution to this problem would be to restrict the systematic analysis of angiosperm megafossils to taxa known from both reproductive and vegetative organs, this approach would greatly restrict information about the flora as a whole, given the dominance of isolated vegetative organs in the megafossil record." The above statement is from page 3 of Upchurch and Dilcher (1990), Cenomanian Angiosperm Leaf Megafossils, Dakota Formation, Rose Creek Locality, Jefferson County, Southeastern Nebraska, U. Separate columns are devoted to extant and extinct taxa.

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