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The clade probably first appeared during Triassic times, possibly as a result of the re-setting of plant evolutionary history following the devastating global extinction event of the Permian Triassic boundary ..." (4. The fossil dataset used by the Cascales-Miñana team is grossly incomplete. Simply put, paleontologic data are required to calibrate and validate molecular phylogenies (Peterson et al. "The interface of these three subject areas (Figure 1 on Page 778), molecular evolution, evolutionary developmental ('evo-devo') biology, and palaeoecology, is the theme of Molecular Palaeobiology, as it [the approach] uniquely integrates the patterns written in the two historical records, genomic and geological ... Labandeira's findings (2014) might also help disprove the notion of a Hauterivian (Lower Cretaceous) origin of flowering plants (Hughes 1994, Friis et al. Errors in molecular-phylogenetic inference may result from effects of LBA (Barrett and Willis 2001, Magallón 2010, Zhenxiang Xi et al. Paraphyly may be underappreciated (Krassilov 2002, Stuessy 2010) and effects on seed plant evolution attributable to possible HT might cloud our understanding of relationships among basal clades of the angiosperm crown group (Bergthorsson et al. "Darwin himself referred to the 'early origin and diversification of angiosperms' as 'an abominable mystery,' and the origin of the flower- and therefore flowering plants- is still a question ..." (page 86, Pamela S. Soltis 2014) Molecular-phylogenetic analyses by Magallón (page 395, 2010) when calibrated with fossil data and compared with different relaxed-clock methods "... Coevolution between phytophagous insect antagonists and Carboniferous, Permian, and Triassic seed plant hosts at the level of their respective developmental tool kits with focus on selective forces that drive the logic of transcriptional regulation is proposed to explain the origin of angiosperms and certain clades of holometabolous insects. Modern syntheses on the abominable mystery of the origin of angiosperms from unknown Paleozoic seed plant ancestors and modern radiations are published by Frohlich and Chase (2007), Maheshwari (2007), Sokolov and Timonin (2007), Zavada (2007), J. After integrating evidence as a whole with our results, the resulting scenario suggests that there is nothing particularly mysterious about the diversification of angiosperms during Cretaceous times or how it is reflected in the fossil record. The preceding statement is an optimistic appraisal of methodology used by Cascales-Miñana et al. Some "current viewpoints" are left out of the analysis. The preceding statement is from page 35 of Armen Takhtajan (1969), Flowering Plants: Origin and Dispersal (translated by C. Conrad Labandeira is apparently less than enthusiastic on the idea of a coevolutionary origin of the group (2014). "Tight coevolution" between animal disperser and plant was probably rare (page 3, Tiffney 2004). 2007) expressed as often disarticulated and shed, wood-, pollen-, seed-, foliar-, and cone- and floral- organs preserved in the fragmentary rock record of the Carboniferous, Permian, and Triassic periods. The image was captured in 1981 while the author was visiting Indiana University. both within and outside the paradigm of transcription-encoding factors ..." (page 129, Niklas 2006). The family Degeneriaceae was discovered in 1942 by I. Endress (1994, 2001 [a book chapter and two papers], 2004), Bateman et al. Several developmental gene families, TFs, and enzymes involved in hormone signaling cascades are known in invertebrates based in part, on experimental studies of the Drosophila model arthropod (S. Wings, halteres, arachnid spinnerets, and insect legs are all organs that develop from limb fields of cells where Ubx expression is prevalent (S. Several insect systematists studying beetle (Coleoptera) evolution are employing some genes and proteins of the insect development tool kit in their phylogenetic analyses (Gómez-Zurita and Galián 2005). The three essays on the succeeding web pages are written from this research perspective. Gómez-Zurita and Galián (2005) discuss the utility of molecular phylogenetic characters appearing in the entomological literature in a review paper, which is organized along the lines of Floyd and Bowman (2007) for land plants (see section below). Understanding the land plant developmental tool kit and gene regulation from a deep time research perspective ties-in with models of cone and floral organization, cell geometry and regulation of growth from SAMs, paleobiology of homeodomain TF trafficking, phyllotaxis, leaf development, and morphogenesis of fertile organs.

Evolutionary-development of arthropod- and plant organs and molecular tool kits is "highly dynamic in evolutionary time" involving the evolution of cis-acting promoters (page 83, Baum 1998). Reviews by Rothwell (1987), Arthur (2002), Meyerowitz (2002), Becker and Theißen (Figure 1, page 468, 2003), Niklas (2006), Rothwell et al. A key paper on the control of insect body size by Nijhout (2003) outlines the molecular mechanisms involving cis-acting TFs and hormones and environmental controls (nutrition and temperature) behind growth and cell division in hemimetabolous and holometabolous insects. Doyle (1978, 1994, 2005, 2006, 2008, 2012), Friis et al. (2008), Berendse and Scheffer (2009), Friedman (2009), Specht and Bartlett (2009), E. A differing proposal by Dahlgren and Clifford (1982) suggests: "The ancestors of the monocotyledons were probably shrublets or subshrubs which by environmental conditions (a pronounced alternation between wet and dry periods) evolved compact underground stems, mainly short or long rhizomes from which herbaceous aerial stems were developed ..." The preceding quotation is from page 344 of R. Class 1c resin constituents associated with Mesozoic angiosperm amber are known from 320 million year old (Paleozoic) amber samples (Bray and K. Almost certainly, accomplished studies of the Amborella trichopoda genome, though useful in disentangling aspects of the evolution of GRNs and horizontal transfer (HT), will not help paleobiologists determine the origin(s) of angiosperms. 2014), and detailed analyses of paleobiological data (Labandeira 2014). If fertile spur shoots are demonstrably ancient organs known from late Paleozoic seed plant fossils then how could the flower possibly originate in the late Mesozoic? "The flower remains ill-defined and its mode (or modes) of origin remain hotly disputed; some definitions and hypotheses of evolutionary relationships preclude a role for the flower in delimiting the angiosperms." The preceding statement is from the abstract on page 3471 of R. Floral morphologies are deeply-conserved in angiosperms according to Melzer et al. According to Stewart and Rothwell (1993) these main steps were: The above bulleted quotes are from pages 461-462 of W. Flowering plants evolved from herbaceous forms possessing ovule and pollen bearing organs that coalesced over time producing modern flowers according to D. Professor Burger proposed six hypothetical trends in the early evolution of angiosperms: The above bulleted quotation is from pages 191-194 of W. Burger (1981), Heresy revived: the monocot theory of angiosperm origin, Evolutionary Theory 5: 189-225. Oleananes, together with ursenes, lupenes, and taraxerenes are important TSBs that belong to a class of Β-amirin triterpenoids (Moldowan and Jacobson 2002). Oleananes occur in fossilized leaf material of certain gigantopterids, bennettitaleans, and flowering plants (Moldowan and Jacobson 2002), but are absent from samples of several other fossil seed plants (D. Problems with Contemporary Data Sets: Controversial assertions abound in the scientific literature of the 20th Century and three categories of credible hypotheses and theories exist (Rothwell et al. None of these ideas when taken as a whole are neither compelling or plausible to many scientists, including the author. Based on a discussion of floral evo-devo by Becker (2016), there are other points of view to be considered. Deciphering evo-devo of short- (spur-) shoots on growing mother plants in hybridizing Permo-carboniferous seed plant populations is probably a central tenet in disentangling at least some aspects of the allopolyploid origin of the flowering plants. Some of the historical syntheses include Arber and Parkin (1907), I. Bailey (1949), Edgar Anderson (1934), Axelrod (1952, 1970), Leppik (1960, 1968), Raven and Kyhos (1965), Cronquist (1968), Thorne (1968), Melville (1969), Takhtajan (1969, 1976, 1991), Raven and Axelrod (1974), Stebbins (1958, 1974), C. Beck (1976), Hughes (1976, 1994), Meeuse (1979), Nair (1979), Krassilov (1977), Retallack and Dilcher (1981 [two papers]), Asama (1982, 1985), Melville (1983), Crane (1985), Meyen (1986, 1988), Dilcher (1986, 2000), J. Doyle and Donoghue (1986, 1987), Endress (1987), Friis et al. If the answer to the preceding question is "yes," how does this evo-devo mechanism affect arthropod antagonist body allometries and population ecology? Further, the evo-devo of flight is yet another conundrum in paleoentomology (Grimaldi and Engel 2005). Poleward migration of early angiosperm flora - angiosperms only displaced the relict Jurassic-type flora at high latitudes in late Cretaceous time. (2017) compile particularly relevant reference lists. Flowering material of Degeneria vitiensis is shown in the right-hand image (photographed by Paddy Ryan, Ph. Fragrance of this species resembles Cananga odorata according to Professor Al Smith (A. While discussing the effects of ice-house/hot house planetary climatic switches on expansion of land plant invertebrate herbivores Labandeira (2006) states: "One possibility is that these atmospheric variables have direct physiologic consequences on the selection and turnover of particular plant clades globally, which in turn elicit an associational response from selected clades of insect herbivores." The preceding statement is quoted from page 425 of C. Labandeira (2006), The four phases of plant-arthropod associations in deep time, Geologica Acta 4(4): 409-438. Additional compilations on the origin of angiosperms and floral morphology include Krassilov (1991), Thorne (1992), Endress (1993, 2001 [a book chapter and two papers], 2004), Friedman (1992 [two papers]), Stewart and Rothwell (1993), Nixon et al. Studies on Drosophila melanogaster eggs, specifically, artificial size-selection experimentation, affects larval patterning and body allometry (Miles et al. Do host seed plant brassinolides and other hormones affect insect antagonist egg size, potentially controlling larval tissue patterning? At the very earliest, flying insects were known from the Devonian Period. The International Journal of Plant Sciences devotes most of Number 7 of Volume 169 (2008) toward the ongoing search for the earliest flowers, based on an international symposium held during the summer of 2007 at the Swedish Museum of Natural History (von Balthazar et al. More than twenty articles in Volume 96, Number 1 of the American Journal of Botany explore the origin, evolution, and radiation of flowering plants to celebrate the Charles Darwin Bicentennial (Stockey et al. Conrad Labandeira's several reviews on fossil insect-plant phytophagous associations (Labandeira 2000, 2006, 2007 [two papers], 2010, 2014) contain extensive bibliographies. 2008) and assembly of chitin and cuticle proteins into the exoskeleton (Charles 2010, Moussian 2010). Another Hox protein Abd-B, when combined with the Dsx enzyme, represses expression of the wg gene in fruit flies (W. I also add hexamerin moulting storage proteins which are related to hemocyanin respiratory enzymes (Burmester et al. 2006, Burmester and Hankein 2007), JH esterases, vitellogenin genes and yolk proteins (Isoe and Hagedorn 2007), pheromone chemoreceptors (Robertson and Wanner 2006), and certain nuclear receptor proteins (Bonneton et al. 2008) including ultraspiracle, and ecdysone inducible TFs to the list of molecular developmental tools among early diverging arthropod lineages. The first appearance of insect wings in the rock record of the Paleozoic Era has yet to be established.