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Doyle (1991, 2000), Frohlich and Parker (2000), Friedman and Floyd (2001), G. The evo-devo research perspective could help us decipher more than 400 million years of insect and seed plant evolution and the enigmatic origins of flowering plants and interacting Holometabola. (2014), and Tomescu (2016), among others, are useful in understanding the developmental systems of animals, fungi, and plants. Several neurosecretory hormones play an important part in mechanisms that regulate cell division and growth including insulin-like peptides (Drosophila insulin-like proteins [DILPs] and bombyxins), chitenase-derived imaginal disk factor proteins, the steroid hormone ecdysone, local autocrine and paracrine TFs, and brain neurosecretory prothoracicotropic hormone (PTTH) (Nijhout 2003). 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. 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 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.

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.

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.

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 in the following essay to explain the origin and evolution of flowering plants and certain Holometabola. A second species of Degeneria has been reported (A. Despite several decades of effort by morphologists, paleobotanists, and plant biologists, the origin of angiosperms remains enigmatic and mysterious. Taylor and Hickey (1996 [a book and one paper]), D. Interestingly, many naturally-occurring plant sesquiterpene esters and lactones are bioactive and exhibit insecticidal properties. Molecular diversification of the Hox gene complex over the course of 600 million years of metazoan evolution is analogous to the 400 million year old molecular evolution of MIKC-type MADS-box genes and related cis-acting TFs of land plants (Theißen et al. Evolution of the Hox complex probably involved small gene duplications, WGDs, divergence of homeodomains, disintegration of the Hox cluster at breakpoints, and rapid changes in the nucleotide sequence of homeodomains (S. Shrub-like lignophytes or small trees produced reproductive modules, which were exploited by flying insects. 2007) and caste polyphenism in holometabolous wasps (J. Understanding the nature and timing of early molecular diversification of homeotic selector genes, developmental proteins, nuclear receptor proteins, and cis-acting TFs of both invertebrate antagonists and vascular plant hosts might be a critical first step in understanding the Paleozoic origin of holometabolous insects and their putative coevolution with the earliest angiosperms.

I discuss potential coevolution of insect and seed plant helix-turn-helix proteins, specifically Engraled and Leafy enzymes that bind to cis-regulatory promoters controlling downstream expression of genes determining paedomorphic insect body patterns and plant cone and floral organ development. (2017) report low support ( The Fiji Islands have long been of interest to biogeographers (Raven and Axelrod 1974, Thorne 1986, Morley 2001), to geologists as a tectonic puzzle (Rodda and Kroenke 1984), and to botanists as a "cradle of flowering plants" (title, Chapter 12, Takhtajan 1969), where some "missing links in the chain of angiosperm phylogeny" are known (page 141, Between Assam and Fiji, Takhtajan 1969). There are several conifers endemic to the Fiji Archipelago including Agathis vitiensis, Acmopyle sahniana, Dacrycarpus imbricatus, Dacrydium nausoriense, Dacrydium nidulum, and Decussocarpus vitiensis. The only known species at the time, Degeneria vitiensis (pictured below), combines a number of primitive features that have ignited many debates (I. Some paleontologists regard the problem of flowering plant origins, "... Juvenile hormone and its homologs are integral in vitellogenesis (Hartfelder 2000), regulation of moult cycles (Truman and Riddiford 2002), and caste development and behavior in social Hymenoptera (Guidugli et al. Were bioactive brassinolides and sesquiterpenes manufactured by Paleozoic seed plants used as chemical warfare agents to affect growth, development, and behaviour of herbivorous insects? Another avenue of deduction somehow ties-in insect evo-devo of wings from gill halteres with increases in atmospheric oxygen during the De CARB. The place and time to begin a molecular phylogenetic analysis is the late Frasnian-Famennian Age hypoxic icehouse that extended into the Tornaisian Age of the Carboniferous Period.

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Erbar (2007) summarizes past ideas on a supposed Mesozoic origin of angiosperms from the research perspective of evolutionary-development (evo-devo). Retallack and Dilcher (1981) presented in-depth discussion of Melville's ideas on a glossopterid ancestry of the angiosperms including a reanalysis of glossopterid fructifications. Others suggest that flowering plants evolved from multiple, unrelated seed plant lineages (Edgar Anderson 1934). 2002) and Nair's Triphyletic Theory (Nair 1979) are best placed in this paragraph. Eichler (1976) proposed that unisexual gymnosperms may be the ancestors of angiosperms. Finally the column labeled "Paraphyly or Polyphyly" denotes whether the scientific paper in question attributes the origin of flowering plants to a natural, intergeneric hybridization event, allopolyploidy, or events that brought together two or more distinct lines of seed plant evolution. Doyle and Donoghue 1986, 1987) and classic research by Arber and Parkin (1907), Edgar Anderson (1934), Axelrod (1952), Ehrlich and Raven (1964), Raven and Kyhos (1965), Takhtajan (1969, 1976), and Raven (1977), dovetail with- and potentially support a coevolutionary hypothesis on the origin of flowering plants, which is developed on the following pages of the web site for purposes of classroom and seminar debate and discussion. Doyle 2008) of perianth parts, microsporophylls, and megasporophylls to form a flower was an improbable and unnecessarily complicated saltational event punctuating a long and gradual evolutionary history of angiosperms. Simply put, massive, shortened bisexual cone axes bearing megasporophylls, laminar microsporophylls, and spirally-arranged foliar tepals, probably existed in populations of poorly understood Paleozoic seed plants described as gigantopteroids and Vojnovskyales, groups omitted by J. Doyle and others in their many published phylogenetic analyses.

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