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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. On the other hand, certain hemimetabolous bugs (Hemiptera) possess abdominal stretch receptors that activate secretion of PTTH (Nijhout 2003). 2005, among others) of the Drosophila Hox complex are: Ancestral arthropods possess two additional homeotic selector genes of the Hox cluster that together comprise the HOM-C, ten gene complex (see discussion in Negre et al. These additional genes are: Genomic analyses suggest that derived winged insects lost functional copies of ftz and Hox3 through disintegration of the HOM-C complex (Negre et al. Duplication of the Hox3 gene of ancestral Cyclorrhaphan flies gave rise to two maternal effect genes, bcd and zen (Stauber et al. Based upon this study it is important to include Hox3 as part of the ancestral diverging insect developmental tool kit. Possible candidates for the early divergent insect developmental tool kit might include certain homeotic selector genes of the Hox complex such as homologs and paralogs of abd-A, Abd-B, Hox3, pb, Scr (Rogers et al. 2002) are probably behind many insect body plan novelties seen in the paleontologic record of the past 400 million years of arthropod and crustacean evolution (Pavlopoulos and Akam 2011, Pavlopoulos and Averof 2002). Plant evolution occurs as variation in genetic and epigenetic developmental processes is winnowed by ecology..." The preceding quotation is from page 161 of P. Once JH circulating in the hemolymph is destroyed by juvenile hormone esterases, then PTTH secretion resumes under circadian (22-24 hour) photoperiodic control (Nijhout 2003). The importance of Ubx protein encoded by the Ubx gene in the early divergent insect developmental tool kit cannot be neglected in the present analysis since significant changes in the carboxy-terminal (C-terminal) region (Galant and Carroll 2002) and serine/threonine phosphorylation sites (Ronshaugen et al. (1985), Labandeira and Sepkoski (1993), Farrell (1998), Labandeira (1998), Danforth and Ascher (1999), Grimaldi (1999), Wilf et al. 2009, Specht and Bartlett 2009, Licausi 2011, Glover 2014, among others). Evidence of arthropod-plant interactions in the Upper Triassic of the Southwestern United States. All total in this rich flora of some 6,000 species, there are 812 endemic angiosperms and conifers, 12 endemic genera, and one endemic flowering plant family (A. Distant pinnacles and spires are weathered calc-alkaline Miocene andesites known as the Namosi Volcanics (Rodda and Kroenke 1984). (2008), Specht and Bartlett (2009), Dilcher (2010), D. Isoptera (termites) are hemimetabolous insects (Grimaldi and Engel 2005). The insect developmental tool kit is comprised of certain homeotic selector genes (including Hox genes), zygotic (gap- maternal-, and pair-rule-) genes, field-specific selector genes, compartment selector genes, cell-type-specific selector genes, and segment polarity genes; and the TFs they encode (Rosenberg et al. In addition, the insect developmental tool kit is comprised of controlling factors behind the cessation of insect growth including bioactive PTTH, JH, juvenile hormone esterases, and ecdysone steroids (Truman and Riddiford 2002, Nijhout 2003, S. Juvenile hormone biosynthesized in the corpora allata of the insect brain is a sesquiterpenoid epoxide methyl ester (Hartfelder 2000). Key elements of the Drosophila molecular tool kit include: Evolution of the Hox complex. Ice-free terrestrial environments in the Late Silurian were covered in vegetation. Understanding the origin and diversification of holometabolous insects in deep-time.
The image to the right is the passive insect trapping flowering plant, Darlingtonia californica (Sarraceniaceae, Ericales, Asteranae), photographed by the author at a seep on Eight Dollar Mountain located in the Klamath Region of western North America. 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. The picture of the rock slab on the left is of an indeterminate pentamerous fossil rosid flower (Celastrales, Rosanae) collected by Professor David L. Three of the largest islands (Viti Levu, Vanua Levu, and Taveuni) support harmonic "continental" floras (A. A common gnetophyte (Gnetum gnemon) and a narrowly distributed cycad (Cycas rumphii) occur in the archipelago. as intractable a mystery today as it was to Darwin 130 years ago" (page 318, Rothwell et al. Simply put, the origin of angiosperms is a conundrum. Another important reason for students of insect-seed plant coevolution to be conversant with arthropod tool kits is that evo-devo of the anterior (head) segment is linked to feeding, pollinating, and sensory perception. According to the discussion in Chapter 6 of Grimaldi and Engel (page 158-159, Insects Take to the Skies, 2005) a "plethora of ideas" on the evo-devo of insect flight "can be distilled into two current but contrasting theories." Studies of pterygote and polyneopteran nymphs suggest that wing pad development evolved independently several times over the past 400 million years (Haug et al. Respiratory enzymes, specifically hemocyanins and hemoglobins, and moulting storage proteins (hexamerins) are key elements of the early divergent arthropod developmental tool kit that tie-in with the evolution of insect legs and wings from bilaterian gills. Interestingly, hexamerins are also implicated as silencers of JH signaling in neotenous castes of hemimetabolous termites (X. Certain details of the Frasnian-famennian boundary extinction (De CARB) are discussed in a later section. New occurrences of the controversial late Triassic plant fossil Sanmiguelia Brown and associated ichnofossils in the Chinle Formation of Arizona and Utah, USA.
Dilcher from the Lower Cretaceous Dakota Formation of North America. Tropical forests of the larger islands yield ten genera of monocotyledonous palms including the monotypic Alsmithia longipes, and the enigmatic magnoliid flowering plant family, Degeneriaceae. Historical Context: Many bibliographies on angiosperm floral diversity and the origin and evolution of flowering plants are available. Labandeira (2010) states: The aforementioned passage is from page 471 of C. Labandeira (2010), The pollination of mid-Mesozoic seed plants and the early history of long-proboscid insects, Annals of the Missouri Botanical Garden 97(4): 469-513. Ancient insect wings probably functioned as respiratory organs. Molecular model systems used as tools in beetle genomic research and phylogenetic studies include proteins central to development (JH esterases), diapause proteins, heat shock proteins, ultraspiracle (an ecdysone nuclear receptor protein), cuticle proteins, hexamerins, genes encoding vitellogenin, and apolipophorins, among others (see review by Gómez-Zurita and Galián 2005). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 268(1): 65-82.
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.