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Were insect and shrub coevolutionary compartments of the late Paleozoic hypoxic icehouse and later hot house, venues of the first angiosperms? This question among others is explored in this first of three essays on the origin of angiosperms. Long-branch attraction (LBA) continues to cloud molecular-phylogenetic studies of seed plants, including angiosperms (Lipeng Zeng et al. Evolutionary-development of early land plants was probably intertwined with regulatory changes in polycomb repressive 2 gene complexes and other stem cell factors as evidenced from studies of the extant model bryophyte Physcomitrella (Okano et al. Floyd and Bowman (2007) are the first workers to estimate the developmental tool kit of early land plants including Paleozoic seed plant homeotic genes potentially important in the later evolution and diversification of angiosperms and origin of the first flowers from bisexual cone axes sensu Melzer et al. The work by Floyd and Bowman (2007) focuses on a molecular-phylogenetic analysis of Chara (a green alga), Physcomitrella (a moss), Selaginella (a lycophyte), Arabidopsis (angiosperm malvid), Antirrhinum (angiosperm asterid), Oryza (angiosperm monocot), Populus (angiosperm fabid), Picea (gymnosperm conifer), and Pinus (among others).

Certain aspects of coevolution of Mesozoic arthropods and seed plants that have a bearing on the origin and diversity of angiosperms are reviewed by Takhtajan (1969), Raven (1977), Thien et al. A review of plant homeobox genes and homeodomain proteins offers additional insight into critical elements of the land plant developmental tool kit (Mukherjee et al. Many developmental gene families and cis-acting TFs have been identified in land plants (Langdale 2008, Mukherjee et al.

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.

Caytoniales and angiosperms diverged from a common ancestor with Bennettitales in the Lower Triassic according to Cascales-Miñana et al. Why assume that flowering plants constitute a single clade first appearing 256 MYA without discussing Mathews (2009), Mathews et al. Discerning Fingerprints of Developmental Regulation: This chapter of the essay considers experimental approaches and paleobiological evidence drawn from the research perspective of evo-devo, which is necessary to identify lineages of seed plants involved in the origin and evolution of flowering plants.

D., and Mark's help at the delnortea beds is gratefully acknowledged. the [angiosperm] clade probably first appeared during Triassic times," which is a stratigraphically-perplexing Gordian Knot. The preceding statement is quoted from page 399 of David Grimaldi and Michael S. This challenging and daunting approach was facilitated by ready access to several world class research libraries at the University of California, Berkeley. The enigmatic Paleozoic plants Spermopteris and Phasmatocycas reconsidered.

Arthropod body allometry is intertwined with development of larval and imaginal disc tissues (Stern and Emlen 1999, Shingleton et al. 1997), Ubx (Pavlopoulos and Akam 2011), and the field-specific selector gene necessary for limb development in Drosophila (Diptera) known as dll (S. Fushi-tarazu protein encoded by the ftz gene, intracellular tertiary enzyme structure folding environments, and the apparent flexibility of Ftz and other Hox proteins in the evolution of arthropods, are discussed in a recent review by Merebet and Hudry (2011). These studies, among others underway or already published by Sean Carroll and colleagues, underscore the importance of Hox proteins in evolution of the arthropod tool kit. Mesozoic paleogeography and early angiosperm history.

Ontogeny is thus the creative force behind botanical diversification, and small modifications at the genetic level may have a disproportionate effect on plant form as their consequences cascade and multiply through development. Kenrick (1997), Diverted development of reproductive organs: a source of morphological innovation in land plants, Plant Systematics and Evolution 206: 161-174. From the research perspectives of insect- and floral biology, and paleoentomology and floral morphology, scaling data might be applied to understanding and computing theoretical morphospace of whole invertebrate and/or plant organs (Jeune et al. Prothoracicotropic hormone and/or ecdysone secretion in Holometabola is negatively controlled by juvenile hormone (JH) (Truman and Riddiford 2002).

Taylor and Hickey (1992, 1996), Loconte (1996), and Krassilov (1997, 2002), among others. "The idea is that plants have a plastic and modular developmental system such that simple changes in regulatory genes need not lead to inviability but can generate novel, potentially favored phenotypes." The preceding quotation is from page 83 of D. "Ontogeny in land plants can be viewed as a complex, partly hierarchical, series of developmental processes, which together with their underlying genetic controls, provide the raw material for morphological innovation. The interface between development and ecology may be studied from such perspectives, among others (Enquist et al. "In theoretical morphospaces, the axes of the reduced space are determined by a small set of parameters of morphogenetic or other biological models, derived from theoretical considerations rather than from the organisms themselves" (page 841, Chartier et al. Scaling studies of reproductive short- (spur-) shoots of living Ginkgo are particularly revealing to plant morphologists (Christianson and 2009). Cessation of growth in holometabolous insects leading to a new moulting cycle is triggered by PTTH that initiates the ecdysone growth regulatory cascade.

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.

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Could paleoecologists benefit by studying experimental, 3-D printed artificial constructs of shoots and protoflowers in theoretical morphospace? By measuring and scaling detached and shed foliar and cone- floral-organs, and by combining these data with studies of permineralizations, "fingerprints of developmental regulation" (quoted from page 723, Sanders et al.

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