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Anatomically preserved lycophytes of the Lycopodiales and Selaginellales have been discovered among a diverse assemblage of plants and fungi in carbonate marine concretions at the Apple Bay locality along the shore of Holbert Inlet near the northern end of Vancouver Island, British Columbia, Canada. Lycopodialean stems are plectostelic and actinostelic, branch dichotomously, and are similar to both Lycopodicaulis oellgaardii and Lycoxylon spp. The Selaginella specimens represent the first anatomically preserved Selaginellales with excellent internal cellular preservation in the fossil record, and are described as Selaginella quatsinoense Rothwell et Stockey sp. nov. Stems have three and five exarch, monarch stelar segments, each of which is surrounded by an aerenchymatous endodermis with trabeculae. The leaf base is indented on the adaxial surface, suggesting the position of a ligule. These fossils document that species with diagnostic internal anatomy of modern Lycopodiales and Selaginellales evolved no later than the Valanginian of the early Cretaceous. [ABSTRACT FROM AUTHOR]

Premise: An anatomically preserved fossil fern sporeling has been discovered in a Lower Cretaceous marine concretion from Vancouver Island, British Columbia, Canada, providing an opportunity to characterize rhizome growth from an extinct species. Methods: The specimen was studied from serial transverse sections prepared by the cellulose acetate peel technique. Results: The rhizome ranges from ~0.7 to 1.1 mm in diameter, has a sclerenchymatous pith, a stele that attains a dictyoxylic architecture, and sclerenchymatous outer cortex, features that are characteristic of osmundaceous rhizomes. Cauline xylem forms a medullated protostele or solenostele at some levels, but is dissected into discrete xylem bundles at others. Fronds diverge in a helical phyllotaxis, range up to 1.1 mm in greatest dimension, and have a C‐shaped trace and outer cortex of sclerotic cells. Inside the sclerenchyma of the petioles are two lateral sclerotic strands and a sclerotic bundle adaxial to the trace. Together, these characters reveal the occurrence of a new species, Todea minutacaulis and provide evidence for developmental changes that occur in the rhizome as the sporeling increased in size. Conclusions: Small size of the specimen, medullated protostelic‐dictyoxylic solenostelar vascular architecture, and incompletely sclerified cells apically reveal that osmundaceous sporeling development has remained constant since at least the Early Cretaceous. Together with Todea tidwellii and Osmunda vancouverensis that also are present in the Apple Bay flora, this sporeling documents the occurrence of a multispecies assemblage of osmundaceous ferns and demonstrates that the genus Todea was diversifying rapidly by the Early Cretaceous. [ABSTRACT FROM AUTHOR]

Premise of research. Two permineralized inflorescences have been found in calcium carbonate concretions from the Eocene Appian Way locality on Vancouver Island, British Columbia, increasing sampling density and helping clarify the pattern of phylogeny for Alismatalean monocots. Methodology. Serial sections of specimens were prepared by the cellulose acetate peel technique and studied with light microscopy. Phylogenetic analyses using a morphological matrix of living and fossil Araceae and Acoraceae were performed using TNT version 1.5 to help establish relationships of the fossil inflorescences. Pivotal results. Inflorescences are cylindrical spikes, >37 mm long and 8.1 mm wide, with a peduncle at least 14.5 mm long, bearing numerous helically arranged, bisexual sessile flowers with hooded tepals. Fruits are trilocular, with basal axile placentation and one seed per locule. Remnants of probable stamens are present in most flowers. Conclusions. Flower structure was compared to those of other petalloid, spicate, bisexual flowers in the monocot families Acoraceae and Araceae (Alismatales). The number of carpels is similar to that in Acorus (Acoraceae) and Spathiphyllum (Araceae, Monsteroideae) and some Pothoideae. Single-seeded ovaries are common in Araceae, while the locules in Acorus contain several apically attached seeds. Phylogenetic analyses place Appianospadix in Proto-Araceae near the base of the aroid tree. This inflorescence with a unique combination of characters is described as Appianospadix bogneri , a new genus and species of Alismatales (family: Araceae). [ABSTRACT FROM AUTHOR]

Anatomically preserved fossil gleicheniaceous fern remains in carbonate marine concretions from Vancouver Island, British Columbia, Canada support the development of a whole plant concept for an Eocene species of Gleichenia , and provide data to develop the first organismal concept for an extinct species of Gleichenia from the Cenozoic fossil record. New information for completing the plant reconstruction includes anatomically preserved pinnule fragments with attached radial exindusinate sori of six to seven annulate sporangia containing trilete spores. While the overall pattern of evolution within Gleicheniaceae remains incompletely understood because well-documented fossils from pre-Cretaceous deposits are inconsistently preserved, this organismal plant concept strengthens evidence that all diagnostic features of the family Gleicheniaceae were present, and that essentially modern species were well-established and diversifying during the Cretaceous and Paleogene. • First whole plant reconstruction of a Cenozoic species of Gleichenia. • Characterized by both external morphology and internal anatomy. • Evidence of stems, frond fragments, and sori of sporangia containing spores. • Essentially modern species of Gleicheniaceae evolved no later than the Paleogene. • Family Gleicheniaceae is inconclusively present in pre-Cretaceous deposits. [ABSTRACT FROM AUTHOR]

Premise: Exceptional anatomical preservation of a fossil araucarian seed cone from a marine carbonate concretion from Vancouver Island, British Columbia, Canada provides unusually complete evidence for cone structure including seeds, megagametophytes, microgametophytes, and embryos of an Upper Cretaceous (Campanian) species of Araucaria, providing important new insights into the structure and relationships of Cretaceous Northern Hemisphere Araucariaceae. Methods: The cone was studied from serial thin sections prepared by the coal ball peel technique. Phylogenetic analysis using a modified morphological matrix with both discrete and continuous characters was performed using TNT version 1.5. Results: The nearly spherical cone, 6 × 6 cm in diameter, has helically arranged cone‐scale complexes, consisting of a large bract with an upturned tip and a small, fleshy ovuliferous scale. Vascularization of the cone‐scale complex is single at its origin. Widely winged bracts, with a bulging base, contain numerous vascular bundles, interspersed with transfusion tissue, and a large number of resin canals. Seeds are ovoid, 1.2 cm long, 1.2 cm in diameter. Nucellus is free from the integument, except at its base, with a convoluted apex, containing possible pollen tubes. Megagametophytes and mature cellular embryos occur in several seeds. Conclusions: This small cone with attached, imbricate leaves, wide bracts, and unusually large seeds, most closely resembles those of Araucaria Section Eutacta. Width and continuity of secondary xylem in the cone axis, and intact cone‐scale complexes indicate that this cone probably did not disarticulate readily at maturity. When added to a modified, previously published phylogenetic analysis, Araucaria famii sp. nov. enhances our understanding of the Cretaceous radiation of Northern Hemisphere Araucaria Section Eutacta. [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]

Premise of research. Five permineralized specimens of infructescences and dispersed fruits assignable to the Altingiaceae have been identified from the Late Cretaceous (Coniacian) Eden Main Quarry on Vancouver Island, British Columbia, Canada. Two infructescences and three isolated two-loculed fruits are preserved in calcium carbonate marine concretions. Methodology. Concretions were sectioned using the cellulose acetate peel technique and were included in a phylogenetic analysis employing morphological characters using TNT. Pivotal results. Infructescences are globose ca. 1 cm in diameter with at least 24 two-loculed fruits and a broad central axis. We have been unable to identify any perianth parts, but there are knob-like extrafloral processes similar to the phyllomes of other altingiaceous infructescences. The inner carpel lining consists of a single palisade layer of elongate rectangular cells. The endocarp also contains several layers of thick-walled, elongate sclerenchyma. The mesocarp is well differentiated, with fiber bundles interspersed with parenchyma and resin canals. There are at least 16 seeds (ovules) per carpel with axile placentation, but most of these are immature or aborted. Fruits have septicidal and ventricidal dehiscence. Specimens are assigned to Protoaltingia comoxense Scharfstein, Stockey & Rothwell gen. et. sp. nov. Conclusions. These fossils represent the second-most ancient occurrence of Altingiaceae, and phylogenetic analysis of morphological characters confirms that P. comoxense is most similar to species of Altingia. Results of morphological analyses resolve Altingia and Liquidambar as sister clades, placing P. comoxense as the basal member of the Altingia clade and sister to Altingia spp., thus documenting that evolutionary diversification of the core eudicot family Altingiaceae (Saxifragales) was well underway by the mid–Upper 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]

Three types of anatomically preserved vegetative shoots with features that characterize crown group Equisetwn have been discovered in Lower Cretaceous deposits (≈136 Ma) of British Columbia. Canada, suggesting the genus is much older than currently believed. Specimens include two types of aerial shoots described as E. haukeanum sp. nov. and E. vancouverense sp. nov. and one type of subterranean rhizome. Shoots are 1-2 mm in diameter, jointed, and in cross section have fluted stems with a hollow pith. Distinctive patterns of cortical sclerenchyma and different ridge morphologies characterize each shoot morphotype. Nodes display irregular branching, highly fused leaf sheaths, and a nodal diaphragm. The aerial stem morphospecies have vallecular canals on alternating radii with carinal canals of an equisetostele surrounded by only a few tracheids. No secondary tissues are produced. Bands of surficial stomata flank the furrows of one morphospecies. Rhizomes and aerial shoots are of a similar size, suggesting that the plants were equivalent in stature to the smallest living Equisetuin species. These fossils augment our understanding of evolutionary transformations that led from Paleozoic Archaeocalamitaceae and Calamitaceae to crown group Equisetaceae, suggesting that the initial diversification of Equisetum began far earlier than suggested by molecular-clock-based estimates. [ABSTRACT FROM AUTHOR]

A remarkably diverse Lower Cretaceous (Valanginian-Hauterivian) flora at Apple Bay, Vancouver Island, preserves seed plants at an important time of floristic evolutionary transition, about the same time as the earliest flowering plant megafossils. The fossils are permineralized in carbonate concretions and include tetrahedral seeds within cupule- or carpel-like structures. These enclosing structures, composed of elongate sclerenchyma cells with spiral thickenings that grade externally to a few layers of parenchyma, are vascularized by one collateral vascular bundle and lack trichornes. They apparently broke open to release the tightly enclosed seeds by valves. Seeds are similar to those of the Triassic seed fern Petriellaea, but arc about 100 million years younger and differ in size, vascularization, integumentary anatomy, seed attachment, and number of seeds/cupule. These new seeds ale described as Doylea zetrahedraspernia gen. et sp. nov., tentatively assigned to Corystospermales. Inverted cupules are reminiscent of an outer angiosperm integument rather than a carpel. Like fruits, cupules opened to release seeds at maturity, thereby foretelling several aspects of angiospermy. They show that nearly total ovule enclosure, a level of organization approaching angiospermy, was achieved by advanced seed ferns during the Mesozoic. [ABSTRACT FROM AUTHOR]

The first fossil evidence for the fern genus Todea has been recovered from the Lower Cretaceous of British Columbia, Canada, providing paleontological data to strengthen hypotheses regarding patterns of evolution and phylogeny within Osmundaceae. The fossil consists of a branching rhizome, adventitious roots, and leaf bases. The dictyoxylic stem has up to eight xylem bundles around a sclerenchymatous pith. Leaf traces diverge from cauline bundles in a typical osmundaceous pattern and leaf bases display a sheath of sclerenchyma around a C-shaped xylem trace with 2-8 protoxylem strands. Within the adaxial concavity of each leaf trace, a single sclerenchyma bundle becomes C-shaped as it enters the cortex. The sclerotic cortex is heterogeneous with an indistinct outer margin. The discovery of Todea tidwellii sp. nov. reveals that the genus Todea evolved by the Lower Cretaceous. A phylogenetic analysis combining morphological characters of living and extinct species with a previously published nucleotide sequence matrix confirms the taxonomic placement of T. tidwellii. Results also support the hypothesis that Osmunda s.l. represents a paraphyletic assemblage and that living species be segregated into two genera, Osmunda and Osmundastrum. Fossil evidence confirms that Osmundaceae originated in the Southern Hemisphere during the Permian, underwent rapid diversification, and species extended around the world during the Triassic. Crown group Osmundaceae originated by the Late Triassic, with living species appearing by the Late Cretaceous. [ABSTRACT FROM AUTHOR]

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous. [ABSTRACT FROM AUTHOR]

Reports on fossil ectomycorrhizae from the British Columbia. Effects of hormones produced by living ectomycorrhizal fungi; Evolution of mycorrhizal associations; Features of the Princeton chert fossils; Characterizations of the mycorrhizal rootlets.

Background and Aims: Widespread and diverse in modern ecosystems, mosses are rare in the fossil record, especially in pre-Cenozoic rocks. Furthermore, most pre-Cenozoic mosses are known from compression fossils, which lack detailed anatomical information. When preserved, anatomy significantly improves resolution in the systematic placement of fossils. Lower Cretaceous (Valanginian) deposits on Vancouver Island (British Columbia, Canada) contain a diverse anatomically preserved flora including numerous bryophytes, many of which have yet to be characterized. Among them is the grimmiaceous moss described here., Methods: One fossil moss gametophyte preserved in a carbonate concretion was studied in serial sections prepared using the cellulose acetate peel technique., Key Results: Tricarinella crassiphylla gen. et sp. nov. is a moss with tristichous phyllotaxis and strongly keeled leaves. The combination of an acrocarpous condition (inferred based on a series of morphological features), a central conducting strand, a homogeneous leaf costa and a lamina with bistratose portions and sinuous cells, and multicellular gemmae, supports placement of Tricarinella in family Grimmiaceae. Tricarinella is similar to Grimmia, a genus that exhibits broad morphological variability. However, tristichous phyllotaxis and especially the lamina, bistratose at the base but not in distal portions of the leaf, set Tricarinella apart as a distinct genus., Conclusions: Tricarinella crassiphylla marks the oldest record for both family Grimmiaceae and sub-class Dicranidae, providing a hard minimum age (136 million years) for these groups. The fact that this fossil could be placed in an extant family, despite a diminutive size, emphasizes the considerable resolving power of anatomically preserved bryophyte fossils, even when recovered from allochthonous assemblages of marine sediments, such as the Apple Bay flora. Discovery of Tricarinella re-emphasizes the importance of paleobotanical studies as the only approach allowing access to a significant segment of biodiversity, the extinct biodiversity, which is unattainable by other means of investigation.

Premise of the Study: Mosses, very diverse in modern ecosystems, are currently underrepresented in the fossil record. For the pre-Cenozoic, fossil mosses are known almost exclusively from compression fossils, while anatomical preservation, which is much more taxonomically informative, is rare. The Lower Cretaceous of Vancouver Island (British Columbia, Canada) hosts a diverse anatomically preserved flora at Apple Bay. While the vascular plant component of the Apple Bay flora has received much attention, the numerous bryophytes identified at the locality have yet to be characterized., Methods: Fossil moss gametophytes in more than 20 carbonate concretions collected from the Apple Bay locality on Vancouver Island were studied in serial sections prepared using the cellulose acetate peel technique., Key Results: We describe Tricosta plicata gen. et sp. nov., a pleurocarpous moss with much-branched gametophytes, tricostate plicate leaves, rhizoid-bearing bases, and delicate gametangia (antheridia and archegonia) borne on specialized branches. A new family of hypnanaean mosses, Tricostaceae fam. nov., is recognized based on the novel combination of characters of T. plicata., Conclusions: Tricosta plicata reveals pleurocarpous moss diversity unaccounted for in extant floras. This new moss adds the first bryophyte component to an already diverse assemblage of vascular plants described from the Early Cretaceous at Apple Bay and, as the oldest representative of the Hypnanae, provides a hard minimum age for the group (136 Ma)., (© 2015 Botanical Society of America.)

Unlabelled: •, Premise of the Study: Two ovulate conifer cones, one of which is attached terminally to a short leafy shoot, reveal the presence of a new species of Hughmillerites in the Early Cretaceous Apple Bay flora of Vancouver Island, British Columbia, Canada. This ancient conifer expands the diversity of Cupressaceae in the Mesozoic and reveals details about the evolution of Subfamily: Cunninghamioideae.•, Methods: Specimens were studied from anatomical sections prepared using the cellulose acetate peel technique.•, Key Results: Vegetative shoots have helically arranged leaves that are Cunninghamia-like. Seed cones have many helically arranged bract/scale complexes in which the bract is larger than the ovuliferous scale. Each ovuliferous scale has three free tips that separate from the bract immediately distal to an inverted seed. Several ovuliferous scales show interseminal ridges between seeds.•, Conclusions: This study documents a new extinct species of cunninghamioid conifers, Hughmillerites vancouverensis, expanding the record of the genus from the Late Jurassic to the Early Cretaceous. This new extinct species emphasizes the important role that conifers from subfamily Cunninghamioideae played in the initial evolutionary radiation of Cupressaceae. In light of recent findings in conifer regulatory genetics, we use H. vancouverensis to hypothesize that variations of expression in certain gene homologues played an important role in the evolution of the cupressaceous ovuliferous scale., (© 2014 Botanical Society of America, Inc.)

An anatomically preserved fossil fern rhizome with diverging stipe bases and root traces is described from the Lower Cretaceous (Valanginian to Hauterivian) Apple Bay locality, Vancouver Island, British Columbia, Canada. The specimen is assignable to Solenostelopteris Kershaw, a morphogenus with six previously described species. The Apple Bay fossil is 1.3-1.6 mm in diameter, with parenchymatous pith and cortex, and is described as S. skogiae sp. nov. The xylem of the solenostele is exarch and one to six cells thick. Successive stipes diverge from only one side of the rhizome, implying a dorsi-ventral symmetry and prostrate habit. No trichomes or scales are produced. Diarch root traces emerge from all sides of the rhizome, some associated with leaf trace divergence. The pith and cortex are made up of uniform, thin-walled cells. The Apple Bay rhizome is most similar to S. nipanica Vishnu-Mittre from the Lower Cretaceous Nipania Flora, India, but differs in size and in distinctive tissue zonation in the cortex. This new species is the youngest record of the genus Solenostelopteris in North America, and it emphasizes that both new specimens of fossils and more complete descriptions of living ferns are needed to help clarify our concepts of Mesozoic ferns.