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New York: The New York Academy of Sciences, 203 pp. The paleobiology of insect flight and seed plant mutualisms is also unexplained. 2007) suggest that WGDs were important in the early evolution of the angiosperm stem group. (1984) to pigeon-hole the youngest Permian plant megafossils known at that time. Yao 1983), suggests that gigantopteroids are gymnospermous (T. Little is known of the whole plant morphology of Carboniferous and Permian gigantopterids although some paleobotanists have deduced a vine habit for these gymnosperms incertae cedis (H. Additional discoveries of permineralized sexual structures are needed to better understand the evolutionary position of these seed plants in relation to other anthophytes (including monocotyledonous flowering plants) and older vojnovskyaleans 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. Reproductive structures were borne on modified leaves, clustered in flower-like strobili. Cycadeoidaceae studied by Wieland and Delevoryas include Cycadeoidea, fossilized remains of squat, shrub-like trunks bearing leaves, cones, and pollen recovered from Cretaceous rocks of North America and Monanthesia (see Delevoryas 1962).

Mousseau (eds.), Annals of the New York Academy of Sciences, Volume 1133 Issue, The Year in Evolutionary Biology 2008. Hermaphroditic shoot apical meristems (SAMs) are the focus of the leading model of cone and floral organization (Melzer et al. Further, the evo-devo of flight is yet another conundrum in paleobiology (Grimaldi and Engel 2005). The term "gigantopteroid" was adopted by Mamay et al. The image was captured on film the day the rock slab was unearthed and collected from the Lower Permian (Cisuralian [Leonardian]) Cathedral Mountain Formation. Better preserved fossilized material and more anatomical data are needed to better understand the anatomy and morphology of whole plants in relation to Vojnovskyales and the angiosperms Joinvillea and Veratrumovuliferous 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, bitegmic ovules (Cornet 1986, 1989) The reproductive biology of Sanmiguelia is not clearly understood, but some research progress has been made (Cornet 1986, 1989). These shrubs were cycad or palm-like in overall aspect. Paleobotanists Crepet, Rothwell, and Stockey review the anatomy and morphology of the Bennettitales (Rothwell et al. The book chapter by Crepet and Stevenson (2010) is a critical library resource on Bennettitales. There are two or three main groups of cycadeoids often treated at the family level in the taxonomic hierarchy.

Rudall (2006), Morphological and molecular phylogenetic context of the angiosperms: contrasting the 'top-down' and 'bottom-up' approaches used to infer the likely characteristics of the first flowers, Journal of Experimental Botany 57(13): 3471-3503. Flowering plants did not appear "suddenly," and sophomorical concepts of a so-called "first flower" should be rejected, in my opinion. Labandeira (2000) identifies these (and other) insect-plant mutualisms as "Assemblage 3: Seed Plants and the Earlier Phase of Modern Insect Fauna," which are based on his review of paleontologic evidence from insect mouthparts and gut contents, and seed plant reproductive organs. Taylor 1999), stems possess a bifacial cambium with vessels (H. Images are 280 million year old permineralizations photographed by the author in 1982 a couple days after the fossils were excavated from the bedding plane of Unit 5 of Section IV of the uppermost Cathedral Mountain Formation, Del Norte Mountains, North America. Paleozoic spermopterids are a relatively unknown group. Were ovule-bearing leaves of Sobernheimia jonkeri parts of enormous bisexual gigantopteroid proanthostrobili with microsporophylls and perianth segments shed and therefore, evading understanding of whole reproductive axes? (2014) Schachat and coworkers report two "overwhelmingly herbivorized taxa," Auritifolia waggoneri (Herbivory Index 3.08) and Taeniopteris sp. Interestingly, "conspicuousness" of certain foliage was suggested as a cue for selective paleoherbivory in these plants (Abstract, Schachat et al. Results of this study beg at least two questions, among others. Eretmonia microsporophyll, redrawn from Surange and Chandra (1975, text-fig. " (page 321, Discussion, Fertilization in Welwitschia, Ephedra, and Gnetum, Friedman 2015). This begs a question, what was the mode of reproductive development and fertilization in gnetalean seed plants of the Permo-carboniferous (Z.-Q. (2002), Mundry and Stützel (2004), Won and Renner (2005), Hajibabaei et al. Intriguing fossilized evidence in Paleozoic rocks of whole plant organs of Sergeia neuburgii (Vojnovskyaceae, Vojnovskyales) is published on page 1072 of Rothwell et al. Unfortunately these fossils cannot be easily screened for invertebrate remains possibly intercalated between the helically arranged leaf permineralizations, because the fossil was transported with other turbidites some distance from the shoreline (Rothwell et al. The image to the left consists of four drawings showing the fructification morphology of certain vojnovskyaleans. The locality of Chekarda-1, the Kungurian, Lower Permian of the Middle Fore-Urals.

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. The resurgence of terrestrial animal and plant life recovering from the PT probably led to evolution of the first seed plant pollinators (Cornet 1989). Taylor 1998, 1999); additional paleontologic data are needed to reconstruct whole plants and nodal anatomyinadequate data, possibly phyllospermous (X. Attachment details of leaves to stems are unclear (Z.-Q. The fossil I collected is the only known microsporophyll of a gigantopteroid, and is left without a complete diagnosis of the organ and the mother plant. It is unknown whether megasporophylls and microsporophylls of Lonesomia mexicana or Phasmatocycas were attached to a massive bisexual cone axis of the same plant or to separate female and male individuals. Herbivorized taeniopteroid foliar organs have been catalogued by Schachat et al. 1d), note that details of sporophyll attachment and sporophyll orientation are uncertain, see text for discussion; × 1. Glossopteris sastrii leaves borne on a shoot, based on Pant and Singh (1974, text fig. However, a seismic shift in favored phylogenetic hypotheses for seed plants associated with the transition from morphological cladistic analyses to DNA sequence-based analyses indicated (and continues to indicate-see above) that Gnetales are not closely related to angiosperms, even if their true phylogenetic affinities remain opaque ... Several groups of vojnovskyaleans had architectural adaptations possibly exploited by Paleozoic insects. Naugolnykh (2001), Morphology and systematics of representatives of Vojnovskyales, Paleontological Journal 35(5): 545-556, reprinted with written permission of Pleiades Publishing, Inc. Naugolnykh and the Paleontological Journal for this contribution. Fructification morphology of representatives of Vojnovskyales: (a, c, d) Paravojnovskya (al. imbricata on a fertile shoot, in axils of scale-form bracts of Nephropsis (Sulcinephropsis).

In turn, PTTH triggers the ecdysone cascade and the larva develops into the pupa, and so forth (Truman and Riddiford 2002). Namely, that some of the taeniopteroid foliage preserved with gigantopteroid leaves in specific sedimentary layers could have been shed from Ginkgo-like short- (spur-) shoots subtended by Cathaysiopteris or Zeilleropteris megaphylls of long-shoots of a completely new kind of seed plant. (1996) that summarizes research on the Permian coal floras of Jiangxi Province, China. A relatively recent report of reproductive structures found preserved in a bedding plane in close association with gigantopterid leaves only adds to the mystery of these plants and their gymnosperm associates (Mei et al. Permineralized gigantopterid foliage and stems belonging to Aculeovinea yunguiensis, Gigantonoclea guizhouensis, and Vasovinea tianii (H. Plumstead (1973) presents an illustrated discussion of the Glossopteris flora, the paleogeography of Gondwana, and Wegener's Theory of Continental Drift. 2007, Xin Wang 2010) and Xingxueanthus sinensis, an angiosperm-like inflorescence from the Middle Jurassic of Asia (Xin Wang and S. 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. Reproductive organs of the shrubs that produced these shed vegetative axes are unknown. Nilssoniopteris vittata, redrawn from Harris (1969, fig. vittata, detail of stoma showing paracytic guard cells and sinuous anticlinal flanges, based on Harris (1969, fig. A pollen chamber is absent and the micropyle is closed by a nucellar plug (Rothwell et al. even if the genus is illegitimate, the various species are validly published (if done according to the rules of that time) and can, therefore, be legitimate ...

In Manduca sexta (Lepidoptera), PTTH is released by the insect brain to initiate a moult. Yet, there is another possible but unexplored interpretation to be proposed that opens windows to greater morphological, paleobiological, and taphonomic precision. Permian Gigantopteris of China unearthed and studied by X.-G. Yao in 1983, yield a rare glimpse of fertile material: ovules and pollen-bearing organs were attached to leaves and leaf-midribs, but the anatomy and placement of the connections is indeterminate. Scutum), compound ovulate structures (Lidgettonia etc.), detached seeds (Pterygospermum and Stephanostoma), fossil leaves (Belemopteris, Gangamopteris, Glossopteris, and Rhabtotaenia, among others), and underground parts (Vertebraria). 1997), Furcula (Harris 1932), Irania (with both androclades and gynoclades), Leptostrobus (a czekanowskialean with stigmatic cupules), Pannaulika triassica (Cornet 1993), and Raunsgaardispermum lusitanicum (palynomorphs resembling Bennettitales and Gnetales) (Mendes et al. Some recent discoveries include Schmeissneria, an intriguing seed plant incertae cedis from the earliest Jurassic Era (Lias interval) of Europe and Middle Jurassic of Asia (Xin Wang et al. Petriellaleans are a group of early Mesozoic gymnosperms with possible evolutionary ties to the Caytoniales (Figures 15.48 - 15.51, and pages 637-639, Chapter 15, Mesozoic Seed Ferns, T. Recent hypotheses propose that the earliest angiosperms may have been small, woody shrubs that colonized disturbed sites in the damp understory of humid forests ... (2014) published a significant find of stems and leaves of Rochipteris alexandriana and permineralized stems of Rudixylon serbetianum (Petriellales) from the Transantarctic Mountains. The nucellus is free from the integument and a tubular micropyle is absent. in April 2014, we all came to the conclusion that the best and simplest option is to leave Taeniopteris as an illegitimate genus and not to conserve it at all ... Despite pivotal discussions on Caytoniales and problematic ideas on origin of the angiosperm carpel (Thomas 1925, Harris 1940, Harris 1951, J.

Ecdysone steroids play a central role in regulation of growth and development of moulting insects comprising the Holometabola (Grimaldi and Engel 2005). Are ovules on some of the taeniopteroid leaves cycad-like or referable to the reproductive structures of Phasmatocycas? Fossilized connections of the leaf impressions with whole plants and reproductive structures of gigantopterids are exceedingly rare or undescribed. Several morphotype genera of glossopterids are representative of detached fossil leaves including Eretmonia, Glossotheca, and Kendostrobus, among others, or isolated pollen sacs (Arberiella, Lithangium, and Polytheca), ovule-bearing leaves (e.g. Key articles on glossopterids are published by Rigby (1967), Surange and Maheshwari (1970), Delevoryas and Gould (1971), Maheshwari (1972), Rigby (1972), Surange and Chandra (1972, 1973, 1975), Holmes (1973), Delevoryas and Person (1975), Chandra and Surange (1976), Schopf (1976), Pant and Choudhury (1977), Gould and Delevoryas (1977), M. White (1978), Rigby (1978), Pant and Nautiyal (1984), Pant and Nautiyal (1984), Pant (1987), and Pigg et al. More recent works are Pigg (1990), Pigg and Taylor (1990), Mc Loughlin (1990), Rigby and Chandra (1990), Pigg and Trivett (1994), Berthelin et al. These poorly known fossil forms include Baisia (Krassilov 1997), Cycandra (pollen cones resembling fossil cycads or Nilssoniales), cupules of Dirhopalostachys (reminiscent of basal flowering plants such as Lesqueria elocata), Doylea tetrahedrasperma (a corystospermous cupule or carpel-like organ, Stockey and Rothwell 2009), Fraxinopsis (Axsmith et al. Seed pods described as Petriellaea triangulata might be homologous with angiosperm carpels but the gymnosperms that shed these organs are as yet unknown. Many authors have noted the similarity of petriellalean cupules to those of the Caytoniales, a group of gymnosperms that continues to figure prominently in theories about the mysterious origin of flowering plants ... coronata, surface of immature 'gynoecium' showing micropyles and interseminal scales, based on Harris (1969, fig. Ovules are born on the distal ends of megasporophylls.

Shrub-like lignophytes or small trees produced reproductive modules, which were exploited by flying insects. (2010), Gorelick and Olson (2011), Marler and Niklas (2011), Olson and Gorelick (2011), and Condamine et al. Cycads have figured prominently in classic papers on the origin of angiosperms (Arber and Parkin 1907, Edgar Anderson 1934, Crane 1985). Co-occurring foliar organs referable to Taeniopteris sp. Contemporary work includes Bose (1968), Delevoryas (1968), Harris (1969), Crepet and Delevoryas (1972), Crepet (1972, 1974), T. Taylor (1973), Harris (1974), Sharma (1974, 1976, 1977), Crane (1986), Pedersen et al.

A review of the floral tool kit by Becker (2016) offers other interpretations of the data. The reproductive SAMs of these Permo-carboniferous seed plants were equivalent to theoretical constructs of the protoflower proposed by Leppik (1960, 1968). The early fossil history of the main taxonomic orders of arthropod antagonists of seed plants presented in this essay is distilled from data published in Rasnitsyn (2002), Grimaldi and Engel (2005), and Labandeira (2006). 1992) bears resemblance to both gigantopteroids and taeniopteroids. The taeniopterid microsporophyll shown to the right (if the rice-grained lumps on the permineralization are proven to be pollen-containing sacs) could be the developmental product of MIKC-type MADS-box B genes such as expression of AP3 or PI homologs along the lines suggested by D. In 1983 Hans Kerp described Sobernheimia jonkeri (page 713, Figure 17.23, T. 1-3, page 232, Weber 1997), which is a gigantopterid according to Professor Weber. "Although we are considering Taeniopteris as a single species, it likely represents multiple species at Colwell Creek Pond [CCP]" (page 856, Geologic and Biologic Setting, The CCP Flora and Comparisons to Relevant Early Permian Floras, Schachat et al. Instead of suggesting that one of the taeniopteroid morphotypes found in these red beds at Colwell Creek Pond [CCP] is Taeniopteris sp. ["the probable cycadophyte" quoted from page 856] (Table 1 on page 858, Schachat et al. Taylor (1993), Iannuzzi (2000), and Pigg and Nishida (2006) compile particularly complete bibliographies on the fossil history of glossopterids. 10E-G), orientation of megasporophyll based on Pant and Nautiyal (1984); × 1. Lidgettonia africana megasporophyll, based on Thomas (1958), Surange and Chandra (1975, text-fig. Some of the research on the anatomy, ecology, evo-devo, fossil history, and molecular systematics of gnetophytes is published by Van Konijnenburg-Van Cittert (1992), Osborn et al. Permineralizations are needed to better understand the anatomy of short shootsglobose head-like bisexual strobilus (e.g. Were the ovules and scales of this plant filled with antiherbivory poisons such as polyacetylenes and toxic cycasins?

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. Data summarized in Table 3 show the extinction of the Palaeodictyoptera and Caloneurida coupled with the appearance of the five new insect orders: Emboidea (appearing in the early Cretaceous Period), Thysanoptera, Hymenoptera (ants, bees, and wasps), Diptera (flies, Figure 306, page 230-231, Rasnitsyn 2002), and Lepidoptera. These specimens and others are deposited in the USNM (Mamay et al. The fossilized leaf imaged to the right is actual size. In Phasmatocycas bridwellii, ovules located on the lower (abaxial) surface of leaves were attached by stalks to leaf midribs but not to the leaf edges as suggested by Cridland and Morris (1960). To the left is the distal portion of an undescribed, retuse Phasmatocycas ovulate leaf that bears resemblance to sterile taeniopterid leaves of Lonesomia mexicana (Plate 3, Figs. 2014), but Evolsonia texana (Herbivory Index 0.95) in these same bedding planes was evidently subjected to less predation than the other two species (Table 1 on page 858, Schachat et al. If detached herbivorized foliar organs provisionally classified as Taeniopteris sp. were shed from short shoots subtended by Evolsonia texana leaves, were not lateral spur shoots of the gigantopteroid seed plant the conspicuous organ being eaten and potentially used by arthropods as habitat? Ottokaria megasporophyll and associated leaf, redrawn from Pant (1977a, fig. Wang 2004), including the MRCA, which was postulated in Figure 3A representing one result of Sarah Mathews concatenated DNA analyses (page 231, 2009)? Scale bar is 1 cm." possibly shrub-like (Mamay 1975, 1976) with scaly, bracetose, and leafy short shoots often fertile in the developing SAMs (Krassilov 1997). 18); × 2." Did sauropods and smaller herbivorous reptiles feed on these fleshy Vardekloeftia seed heads?

If the flowering plant crown node was centered in the Middle Triassic (C. The chapter is split into two sections: the paleobotany of gymnospermous seed plants and a second on the paleoentomology of insect groups which might have ate, bred, and sheltered inside Paleozoic and early Mesozoic vegetation compartments. Ghost lineages are illustrated in the table by a white space with a question [? Based on scanty paleontologic evidence the basal arthropod orders Collembola (springtails), Archaeognatha (bristletails), and Zygentoma (firebrats or silverfish) probably co-occurred with the earliest divisions of vascular plants (Rhyniophyta, Trimerophytophyta, and Zosterophyllophyta; Table 1, page 350, Labandeira 1998; Table 1, page 415, Labandeira 2006). Taylor 1998, 1999), remarkably similar to characters found in bennettitaleans, gnetophytes, and certain extant flowering plants. 2003) detached pieces of a large bisexual cone axis, i.e. The preceding passage is quoted from page 361 of D. Are two morphologically different but sympatric species found in these beds, or does the dimorphic foliage preserved in these rocks belong to an undescribed Permian gigantopteroid seed plant, which is neither a Auritifolia waggoneri or Supaia thinnfeldioides peltasperm, and not a gnetophyte, pteridosperm, or taeniopteroid cycadophyte? Was Eophyllogonium cathayense attached to a gigantopteroid plant with dimorphic, heteroblastic leaves? 2009) might also be detached megasporophylls of a massive bisexual gigantopteroid proanthostrobilus with the microsporophylls and perianth also shed and therefore, evading a complete diagnosis of the whole fertile SAM. Specific examples are wind-dispersed pollen described as the Pennsylvanian callistophytalean morphogenus Vesicaspora (Figure 14.170 on page 598 of T. Much of the morphology and biology of Mesozoic plants should be considered in the light of this substantial herbivore pressure." The preceding quotation is from page 94 of Tiffney (1992), The role of vertebrate herbivory in the evolution of land plants, The Palaeobotanist 41: 87-97.

I concluded in the first essay that divergence of angiosperms from the most recent common ancestor of seed plants (MRCA) was probably much older than suggested by paleontologic data, which casted considerable doubt on all of J. Doyle's so-called combined molecular- and morphological- seed plant phylogenies. Paleontologic Evidence: I now summarize evidence from a paleontologic research perspective to support of a coevolutionary origin of angiosperms and certain arthropod groups. Integers in each cell of Table 3 represent the number of insect families reported from the fossil record boiled down from discussion in Rasnitsyn (2002) including data on non-insect hexapods from Grimaldi and Engel (2005). ] in a cell denotes gaps in the data due to controversial or uncertain familial taxonomic placement. Simply put, detailed study of polished thin-sections of permineralized fertile gigantopterid plant material is required to justify considerations of the magnitude stated in the two papers published by Glasspool et al. Reproduction in gigantopterids is incompletely known (X. Oleonone triterpanes have been isolated from several gigantopterid leaf specimens, fossilized bennettitalean foliage, pyrolized extant angiosperm plant material, and from leaf compressions of flowering plants (D. This specimen is deposited in the USNM (Mamay et al. Were ovulate Phasmatocycas bridwellii leaves (Axsmith et al. Doyle 2008) of an unknown Paleozoic gymnospermous gigantopterid tree or shrub with the pollen-bearing leaves (microsporophylls) and strap-shaped petaloid organs of the perianth missing? What alternative seed plant shoot configurations are possible in theoretical morphospace, based on taphonomic observations and frequency distribution of detached and shed foliar organs observed in discreet bedding planes from the numerous Lower Permian localities where Evolsonia texana and Taeniopteris co-occur? Seed-bearing taeniopterid leaves with reticulate venation were found in the same bedding plane as sterile gigantopterid leaves assignable to Gigantonoclea acuminatiloba and Gigantopteris dictyophylloides. "Since conifers extend back to the Late Carboniferous, this implies that the line leading to angiosperms goes back this far too - an apparent conflict with the stratigraphic record (Axsmith et al., 1998; Doyle 1998a)." This preceding passage is quoted from page 172 of J. Doyle (2001), Significance of molecular phylogenetic analyses for paleobotanical investigations on the origin of angiosperms, The Palaeobotanist 50: 167-188. Several morphological structures seen in extant species of Ceratophyllum are probably homologous with features of the anatomy and morphology of certain living cycads and conifers, and Upper Paleozoic Callistophytales and Vojnovskyales. 2009) and haustorial pollen tubes of certain extant species of Cycadales (D. The image to the right is a drawing of a detached spherical ovule-bearing Vardekloeftia sulcata head with densely packed ovules and interseminal scales, from a bennettitalean bush. Crane (1985), Phylogenetic analysis of seed plants and the origin of angiosperms, Annals of the Missouri Botanical Garden 72: 716-796, reprinted with permission of the Missouri Botanical Garden and Peter Crane. Morphology of the Vardekloeftia and Bennetticarpus plants. Initially, these forces were met by a limited diversity of genetic lineages of plants, but by the end of the Mesozoic, gymnosperms with efficient vegetative growth and abilities to recover from damage (= angiosperms) had evolved.

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