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Premise of research. Numerous large permineralized fossil seeds from the Jurassic of Haida Gwaii, western Canada, provide evidence for the mode and tempo of cycad evolution. Methodology. Fossil seed specimens are studied from external morphology and serially sectioned by the classic cellulose acetate peel technique to reveal anatomical features of the integument, vascular system, nucellus, and gametophyte. Fossils are compared to living cycad seeds that are also examined from external views, dissections, and anatomical preparations. Pivotal results. The fossil seeds are described as Traskia maahlae gen. et sp. nov. Features of the integument and vascular system reveal that T. maahlae is a stem group cycad that shares seed architecture and mode of germination with living Cycas spp. Conclusions. Traskia maahlae adds to a growing body of paleontological data on stem and crown group cycads. Results suggest that modern pollination and postpollination biology and the two contrasting modes of cycad seed germination evolved during the Mesozoic but that crown group cycad species may not have appeared until the Cenozoic. [ABSTRACT FROM AUTHOR]

Premise of research. A single cylindrical conifer seed cone has been discovered in a calcium carbonate marine concretion from the Upper Cretaceous Eden Main Quarry, dated as early Coniacian (∼89 Ma), 35 km northwest of Courtenay on Vancouver Island, British Columbia, Canada. Methodology. The specimen was sectioned with a rock saw, and serial sections were prepared using the classical cellulose acetate peel technique. Peel sections were mounted on microscope slides. Reconstructions of the cone and seeds were rendered with Avizo. Pivotal results. The cone is somewhat abraded, at least 2.3 cm long and 1 cm wide, with the apex missing and most of the base coalified. The well-preserved midsection shows bract/scale complexes inserted at a steep angle. The bract/scale complex has a central rhomboidal shaft and two lateral winglike extensions. Xylem of the cone axis is robust, with secondary xylem tracheids that have circular-bordered pits. A single circular vascular trace enters the bract/scale complex. Three resin canals enter the base of the scale and divide to form up to eight or nine canals distally. Six overlapping, inverted winged seeds, ∼8 mm long and 1 mm thick, with two large, fleshy wings are borne on the adaxial surface of each scale. The nucellus is free from the endotesta to its base. Megagametophytes and embryos occur in some seeds. Cells of the megagametophyte have granular contents. This cone shows similarities in overall structure and vasculature to those of living Taiwania (Cupressaceae); however, the fossil cone size is generally larger, with six seeds per scale versus two, and seeds are extremely large and fleshy with a unique submicropylar structure. Conclusions. This cone is compared with living and fossil taxa of Cupressaceae and described as Comoxostrobus rossii gen. et sp. nov. (Cupressaceae subfamily Taiwanioideae), revealing that taiwanioid Cupressaceae were much more diverse and widespread during the Cretaceous. [ABSTRACT FROM AUTHOR]

A fossil seed cone with characters that have been hypothesized as transitional to the origin of crown group gnetophytes has been discovered in Lower Cretaceous deposits on Vancouver Island, British Columbia, Canada. This cone, described as Protoephedrites eamesii gen. et sp. nov., provides the first anatomically preserved fossil evidence for the evolution of gnetophyte cones with seeds. The cone is a compound shoot system consisting of a primary axis with nodes that bear bracts and axillary fertile shoots in an opposite/decussate arrangement. The secondary fertile shoot axis produces one or two pairs of diminutive bracteoles and a pair of erect ovules in an opposite/decussate pattern. In contrast to crown group gnetophytes, bracteoles subtend rather than surround and enclose the ovules. Ovules have a single multiseriate integument that is adnate to the nucellus in the basal region and free distally. The micropylar tube is short with a thick integument, but it displays distinctive cells of the inner integumentary epidermis that are characteristic of gnetophyte seeds. Each seed produces a large pollen chamber with a uniseriate wall. Paired erect ovules conform to the pleisomorphic morphology previously hypothesized for Ephedra seed cones, thus supporting the proposal that ancestral gnetophyte seeds are borne terminally on oppositely arranged sporophylls. The combination of gnetophyte synapomorphies and putative pleisomorphic characters displayed by Protoephedrites broadens the known range of morphologies for the most ancient gnetophytes. Detailed comparisons to several species of Bennettitales confirm that there are fundamental structural differences separating the seeds of Gnetales from those of Bennettitales, support the hypothesis that the outer seed envelope evolved within the gnetophyte clade, and suggest that Bennettitales are not as closely related to Gnetales as hypothesized by some authors. [ABSTRACT FROM AUTHOR]

A cylindrical permineralized conifer seed cone has been identified from the Officer Member of the Trowbridge Formation, near Izee, in east-central Oregon. The cone is preserved in a Middle Jurassic (Callovian) marine calcium carbonate concretion, associated with araucarian seed cones, conifer twigs and wood, cycad seeds, fern rachides, and lycopodialean remains and was prepared by the cellulose acetate peel technique. The specimen is abraded, 2.8 cm long and 1.3 cm wide, and consists of a cone axis with helically arranged bract/scale complexes, with a large ovuliferous scale and a broad, flattened bract. The bract/scale trace arises as a cylindrical unit from the cone axis, in which the bract and scale traces are separated by ground tissue. The traces separate immediately in the cortex to form a crescent-shaped bract trace and a horseshoe-shaped scale trace with associated sclerenchyma bands that form two triangular bundles toward the distal end of the scale. Ovuliferous scale tissue covers one inverted seed per scale, forming a pocket that contains the seed. Seeds are attached at the chalaza to the inside of the pocket. Seed integuments have an outer epidermis of isodiametric cells with dark contents; one or two layers of palisade cells, occasionally appearing in I-beam configuration; and several inner layers of randomly arranged cells. The nucellus is adnate to the integument, to near the apex, where it forms a cellular mound of tissue. Cylindrical cones with helically arranged bract/scale complexes, bract and scale free to the base, one seed per scale enclosed in a pocket, and triangular sclerenchyma bands are characteristic of the genus Vararaucaria, which recently has been placed in the extinct conifer family Cheirolepidiaceae. This third species extends the known range of Pararaucaria from the Southern Hemisphere to the Northern Hemisphere and emphasizes that the association of araucarian and pararaucarian conifers extended over a wide geographic area during the Jurassic. [ABSTRACT FROM AUTHOR]

* Premise of the study: Pinaceae and nonpinoid species are sister groups within the conifer clade as inferred from molecular systematic comparisons of living species and therefore should have comparable geological ages. However, the fossil record for the nonpinoid lineage of extant conifer families is Triassic, nearly 100 million years older than the oldest widely accepted Lower Cretaceous record for Pinaceae. An anatomically preserved fossil conifer seed cone described here extends the stratigraphic range of Pinaceae nearly 30 million years, thus reducing the apparent discrepancy between evidence from the fossil record and inferences from systematic studies of living species. * Methods: Material was prepared as serial thin sections by the cellulose acetate peel technique, mounted on microscope slides, and viewed and photographed using transmitted light. * Key results: A large cylindrical cone consisting of bract-scale complexes that diverge from the cone axis in a helical phyllotaxis has bracts and scales that separate from each other in the midregion and are of equal length and of nearly equal width. The cone has two inverted and winged seeds that are attached to the adaxial surface of each cone scale and, thus, represents an early member of the Pinaceae. * Conclusions: Eathiestrobus mackenziei gen. et sp. nov. extends the fossil record for well-documented members of the family Pinaceae from the Lower Cretaceous to the Kimmeridgian Stage of the Upper Jurassic. This species also clarifies the set of characters that are diagnostic for seed cones of Pinaceae and reveals possible plesiomorphic characters for seed cones of the family. [ABSTRACT FROM AUTHOR]

Bennettitales is an extinct group of seed plants with reproductive structures that are similar in some respects to both Gnetales and angiosperms, but systematic relationships among the three clades remain controversial. This study summarizes characters of bennettitalean plants and presents new evidence for the structure of cones and seeds that help clarify relationships of Bennettitales to flowering plants, Gnetales, and other potential angiosperm sister groups. Bennettitales have simple mono- or bisporangiate cones. Seeds are borne terminally on sporophylls. They have a unique structure that includes a nucellus with a solid apex, no pollen chamber, and a single integument, and they are clearly not enclosed by a cupule or other specialized structures. Such features differ substantially from Gnetales, flowering plants, and the seed fern Caytonia, providing no compelling evidence for the origin of the angiospermous camel. Cladistic tests were performed to assess the strength of the "anthophyte hypothesis" and possible relation- ships of Bennettitales, Gnetales, and Caytonia to flowering plants. Our results do not support the anthophyte hypothesis for the origin of angiosperms by a transformation of fertile organs that were already aggregated into a cone or flower-like structure. However, the anthophyte topology of the seed plant tree continues to be supported by morphological analyses of living and extinct taxa. [ABSTRACT FROM AUTHOR]

A single cylindrical seed cone 2 cm long, 1.1 cm wide has been found preserved in a calcium carbonate marine concretion from the Hakobuchi Formation (late Campanian-early Maastrichtian) of Hobetsu, Hokkaido, Japan. The cone, attached to a bent peduncle lacking leaves, has helically arranged bract/scale complexes that arise at right angles from the cone axis in the middle of the cone. The cone axis, ca. 1 mm wide, has a broad cylinder of secondary vascular tissue, and lacks a continuous resin canal system. Bract-scale complexes are laminar, cordate-orbiculate, and upturned distally, consisting primarily of bract tissue with no visible scale tip. The vascular trace to the bract/scale complex originates as a rod that divides laterally into several traces at the level of seed attachment. A single resin canal originates at the base of the bract-scale complex abaxial to the vascular strand, but more distally there are up to ca. 15 large resin canals that form a single row. Two to three inverted winged seeds are attached adaxially near the cone periphery. Cone structure and vascularization are most similar to those in the Cupressaceae, Subfamily Taiwanioideae, differing from living Taiwania cryptomerioides by having up to three seeds/scale rather than two, an abruptly upturned bract tip, in details of bract/scale vasculature, and a cone peduncle lacking leaves. This cone is described as Mukawastrobus satoi Stockey, Nishida and Rothwell. Together with previously described Early to Late Cretaceous taiwanioid seed cones from Mongolia and Hokkaido the new species demonstrates that the taxonomically diagnostic characters of such conifers are as subtle as those of Cretaceous and Cenozoic sequoioid Cupressaceae. This realization emphasizes that evolutionary diversification and turnover among taiwanioid conifers during the Cretaceous and Paleogene are probably far greater than currently recognized. [ABSTRACT FROM AUTHOR]

Conifers of the taxodiaceous grade of Cupressaceae were more diverse and widespread during the Mesozoic than they are today. The earliest diverging subfamily, Cunninghamioideae, only includes a single extant genus, but has at least 10 fossil genera. Here, two additional cunninghamioid genera are characterized on the basis of permineralized seed cones from the Upper Cretaceous of Hokkaido, Japan. These conifers display seed cone characters typical of cunninghamioids; however, they have a mosaic of characters that are not seen in any reported conifer of Cupressaceae. They are, therefore, designated as two new extinct species: Ohanastrobus hokkaidoensis gen. et sp. nov. and Nishidastrobus japonicum gen. et sp. nov. These newly reported conifers expand the taxonomic and morphological diversity of cunninghamioids. The stratigraphic and paleobiogeographic records of cunninghamioids and other fossil Cupressaceae with foliate seed cones indicate they peak in diversity during the Cretaceous. The living genera Taiwania and Cunninghamia appear during the Albian and Campanian, respectively, and maintain a nearly continuous fossil record through to today, while nearly all other extinct genera of Cupressaceae with foliate cones disappear by the close of the Campanian. As more ancient cunninghamioids are recovered, our understanding of macroevolutionary patterns of this once diverse lineage will be further elucidated. [ABSTRACT FROM AUTHOR]