Publications by authors named "Ruth A Stockey"

35 Publications

Cynodontium luthii sp. nov.: a permineralized moss gametophyte from the Late Cretaceous of the North Slope of Alaska.

Am J Bot 2021 03 2;108(3):495-504. Epub 2021 Mar 2.

Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon, 97331, USA.

Premise: Mosses are a major component of Arctic vegetation today, with >500 species known to date. However, the origins of the Arctic moss flora are poorly documented in the fossil record, especially prior to the Pliocene. Here, we present the first anatomically preserved pre-Cenozoic Arctic moss and discuss how the unique biology of bryophytes has facilitated their success in polar environments over geologic time.

Methods: A permineralized fossil moss gametophyte within a block of Late Cretaceous terrestrial limestone, collected along the Colville River on the North Slope of Alaska, was studied in serial sections prepared using the cellulose acetate peel technique.

Results: The moss gametophyte is branched and has leaves with a broad base, narrow blade, and excurrent costa. We describe this fossil as Cynodontium luthii sp. nov., an extinct species of a genus that is known from the High Arctic today. Cynodontium luthii is the oldest evidence of the family Rhabdoweisiaceae (by ≥18 Ma) and reveals that genera of haplolepideous mosses known in the extant Arctic flora also lived in high-latitude temperate deciduous forests during the Late Cretaceous.

Conclusions: The occurrence of C. luthii in Cretaceous sediments, together with a rich Pliocene-to-Holocene fossil record of extant moss genera in the High Arctic, suggests that some moss lineages have exploited their poikilohydric, cold- and desiccation-tolerant physiology to live in the region when it experienced both temperate and freezing climates.
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http://dx.doi.org/10.1002/ajb2.1617DOI Listing
March 2021

Extending the fossil record for foliicolous Dothideomycetes: Bleximothyrium ostiolatum gen. et sp. nov., a unique fly-speck fungus from the Lower Cretaceous of Virginia, USA.

Am J Bot 2021 01;108(1):129-144

Department of Botany, University of British Columbia, Vancouver, BC, V6T 1Z4, Canada.

Premise: Fossils can reveal long-vanished characters that inform inferences about the timing and patterns of diversification of living fungi. Through analyzing well-preserved fossil scutella, shield-like covers of fungal sporocarps, we describe a new taxon of early Dothideomycetes with a combination of characters unknown among extant taxa.

Methods: Macerated clays from the Potomac Group, lower Zone 1, from the Lower Cretaceous (Aptian, 125-113 Ma) of Virginia USA yielded one gymnospermous leaf cuticle colonized by 21 sporocarps of a single fungal morphotype. We inferred a tree from nuclear ribosomal DNA of extant species, and coded morphological characters to evaluate alternative, equally parsimonious placements of the fossil in a molecular constraint tree of extant species.

Results: Bleximothyrium ostiolatum gen. et sp. nov. has an ostiolate scutellum of radiate, dichotomizing hyphae. Unlike otherwise similar extant and fossil taxa, B. ostiolatum has tangled hyphae at its scutellum margin. Scutella of B. ostiolatum are connected to superficial mycelium, to intercalary and lateral appressoria, and to extensive subcuticular "mycélium en palmettes". The gymnospermous host has characters consistent with identity as a non-papillate ginkgophyte or cycad.

Conclusions: Bleximothyrium ostiolatum is the oldest known fossil fly-speck fungus that occurs on plant cuticles and has the radiate, ostiolate scutellum known only from Dothideomycetes. Its combination of characters, its scutellum margin, and mycélium en palmettes are unknown in other extant and fossil species, and Bleximothyrium ostiolatum likely represents a new group of fly-speck fungi that may now be extinct.
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http://dx.doi.org/10.1002/ajb2.1602DOI Listing
January 2021

Diversification of crown group Araucaria: the role of Araucaria famii sp. nov. in the mid-Cretaceous (Campanian) radiation of Araucariaceae in the Northern Hemisphere.

Am J Bot 2020 07 23;107(7):1072-1093. Epub 2020 Jul 23.

Department of Botany and Plant Pathology, Oregon State University, 2082 Cordley Hall, Corvallis, OR, 97330, USA.

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.
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http://dx.doi.org/10.1002/ajb2.1505DOI Listing
July 2020

Evolutionary diversification of taiwanioid conifers: evidence from a new Upper Cretaceous seed cone from Hokkaido, Japan.

J Plant Res 2020 Sep 19;133(5):681-692. Epub 2020 Jul 19.

Department of Botany and Plant Pathology, Oregon State University, Corvallis, OR, 97331, USA.

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.
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http://dx.doi.org/10.1007/s10265-020-01214-yDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7429551PMC
September 2020

A new epiphyllous fly-speck fungus from the Early Cretaceous Potomac Group of Virginia (125-112 Ma): , gen. et sp. nov.

Mycologia 2020 May-Jun;112(3):504-518. Epub 2020 Mar 13.

Department of Botany, University of British Columbia , Vancouver BC, V6T 1Z4, Canada.

Fly-speck fungi reproduce via thyriothecia that consist of sporogenous tissue appressed to cuticle surfaces of plant leaves and covered by a shield-like scutellum. Thyriothecial scutella likely evolved repeatedly in Dothideomycetes (Ascomycota), and their morphology varies by lineage. Fly-speck fungi have an exceptionally good fossil record that begins in the Mesozoic. The interpretation of scutellum characters in fossils may provide insights into origins of Dothideomycetes and help calibrate the timing of ascomycete evolution. From sediments of the Lower Cretaceous (125-112 Ma) Potomac Group of Virginia, from Dutch Gap Canal, lower Zone 1, we found scutella similar to those of extant Aulographaceae (Dothideomycetes), attached to a single piece of dispersed coniferous cuticle. We analyze hyphae and scutellum development among four extant Aulographaceae species for comparison with the fossil. The excellent preservation of fungi on the leaf cuticle surface allows us to infer a developmental sequence for the fossil. Scutellum development begins with coordinated growth of multiple neighboring generator hyphae and continues with hyphae producing two-dimensional pseudomonopodial, dichotomous, radial growth. Asci and ascospores were not found. We coded states for seven morphological characters using direct observations of the fossil and eight extant taxa, and using the literature for 28 others. We inferred a phylogeny using nuclear 18S and 28S rDNA of 36 extant taxa, 34 Dothideomycetes and two Arthoniomycetes. The phylogeny includes newly determined sequences from five species, two from Aulographaceae. With a branch-and-bound search, we inferred the most parsimonious placements of the fossil given the molecular tree topology. The parsimony analysis constrained by the rDNA phylogeny places the fossil taxon among stem lineages near Aulographaceae or among the known living members of Aulographaceae. We describe the fossil morphotype as , gen. et sp. nov. The fossil provides the oldest evidence of morphological characters restricted among extant fungi to Aulographaceae.
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http://dx.doi.org/10.1080/00275514.2020.1718441DOI Listing
March 2020

Resolving the overall pattern of marattialean fern phylogeny.

Am J Bot 2018 08 12;105(8):1304-1314. Epub 2018 Jul 12.

Department of Botany and Plant Pathology, Oregon State University, 2082 Cordley Hall, Corvallis, OR, 97331, USA.

Premise Of The Study: Recent clarification of the distribution of Marattiales through time provides the impetus for "total evidence" phylogenetic analyses of a major fern clade with a rich fossil record. These analyses serve as empirical tests for results from systematic analyses of living species and also of the belief that relationships among living species accurately reflect the overall pattern of phylogeny for clades with an extensive fossil record and a large percentage of extinction.

Methods: Species of living and fossil Marattiaceae are analyzed employing a "total evidence approach" via maximum parsimony. Analyses were conducted using TNT implemented through WinClada.

Key Results: Systematic analyses of living species and of living + extinct species provide roughly concordant topologies for living taxa. However, living species of Marattiales are only one component of a much larger clade with two major subclades. One consists of Psaroniaceae and extends through time to at least the Early Cretaceous. The other consists of Marattiaceae and includes all living species. Various analyses support the generic-level clades of living species from earlier analyses, but the arrangement of such clades varies from analysis to analysis.

Conclusions: Marattiales is a monophyletic group that is extremely common in late Paleozoic and early Mesozoic deposits, with a stem group Psaroniaceae and a crown group Marattiaceae. Because Marattiaceae represents only a small component of overall marattialean diversity, living species alone neither account for evolutionary changes within the clade over time, nor accurately reflect the overall pattern of marattialean fern phylogeny.
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http://dx.doi.org/10.1002/ajb2.1115DOI Listing
August 2018

Grimmiaceae in the Early Cretaceous: Tricarinella crassiphylla gen. et sp. nov. and the value of anatomically preserved bryophytes.

Ann Bot 2018 06;121(7):1275-1286

Department of Biological Sciences, Humboldt State University, Arcata, CA, USA.

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.
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http://dx.doi.org/10.1093/aob/mcy015DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6007789PMC
June 2018

Extending the fossil record of Polytrichaceae: Early Cretaceous gen. et sp. nov., permineralized gametophytes with gemma cups from Vancouver Island.

Am J Bot 2017 Apr 19;104(4):584-597. Epub 2017 Apr 19.

Department of Biological Sciences, Humboldt State University, Arcata, California 95521 USA

Premise Of The Study: Diverse in modern ecosystems, mosses are dramatically underrepresented in the fossil record. Furthermore, most pre-Cenozoic mosses are known only from compression fossils, lacking detailed anatomical information. When preserved, anatomy vastly improves resolution in the systematic placement of fossils. Lower Cretaceous deposits at Apple Bay (Vancouver Island, British Columbia, Canada) contain a diverse anatomically preserved flora that includes numerous bryophytes, many of which have yet to be characterized. Among them is a polytrichaceous moss that is described here.

Methods: Fossil moss gametophytes preserved in four carbonate concretions were studied in serial sections prepared using the cellulose acetate peel technique.

Key Results: We describe gen. et sp. nov., a polytrichaceous moss with terminal gemma cups containing stalked, lenticular gemmae. Leaves with characteristic costal anatomy, differentiated into sheathing base and free lamina and bearing photosynthetic lamellae, along with a conducting strand in the stem, place in family Polytrichaceae. The bistratose leaf lamina with an adaxial layer of mamillose cells, short photosynthetic lamellae restricted to the costa, and presence of gemma cups indicate affinities with basal members of the Polytrichaceae, such as , , and .

Conclusions: enriches the documented moss diversity of an already-diverse Early Cretaceous plant fossil assemblage. This is the third moss described from the Apple Bay plant fossil assemblage and represents the first occurrence of gemma cups in a fossil moss. It is also the oldest unequivocal record of Polytrichaceae, providing a hard minimum age for the group of 136 million years.
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http://dx.doi.org/10.3732/ajb.1700002DOI Listing
April 2017

Cretaceous origin of dogwoods: an anatomically preserved (Cornaceae) fruit from the Campanian of Vancouver Island.

PeerJ 2016 21;4:e2808. Epub 2016 Dec 21.

Department of Botany and Plant Pathology, Oregon State University, Corvallis, OR, United States; Department of Environmental and Plant Biology, Ohio University, Athens, OH, United States.

Background: Cornaceae consists of 58 species, all within the genus . The Cenozoic record of is extensive and well documented. Molecular divergence-time studies suggest that crown-group may have originated by the Late Cretaceous. However, there has been no formal report of from Cretaceous deposits. Here, we characterize a permineralized fossil fruit assignable to subg. from the Upper Cretaceous (Campanian) Shelter Point locality of Vancouver Island, British Columbia, Canada.

Methods: Serial sections of the specimen were made using the cellulose acetate peel technique. Peels were mounted onto microscope slides and studied by light microscopy.

Results: The fossil fruit consists of a tri-locular woody endocarp with dorsal germination valves. The locules are sub-triangular to ellipsoidal in transverse section and are separated by thin septa. Endocarp tissue consists of elongated and isodiametric sclereids and secretory cavities. Internal vascular tissue was not observed, but is interpreted to have been located along the outer periphery of the septa for some length, common in many cornalean taxa. There is one seed in each locule, one of which was found to have endosperm and a dicotyledonous embryo.

Discussion: Woody endocarps with germination valves, without central vascular bundles, and with one seed per locule are characteristic of several families within the order Cornales. The interpreted vascular pattern and presence of secretory cavities indicates that the fossil fruit is assignable to subg. . Comparative analysis suggests that the fossil is most similar to , a species described from the Paleocene of North Dakota. This fossil is the first evidence of crown-group Cornaceae from the Cretaceous and sheds light on both the plesiomorphic fruit characters and the timing of the initial diversification of the family and basal asterid lineage, Cornales.
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http://dx.doi.org/10.7717/peerj.2808DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5180587PMC
December 2016

Anatomically preserved fossil cornalean fruits from the Upper Cretaceous of Hokkaido: Eydeia hokkaidoensis gen. et sp. nov.

Am J Bot 2016 09 2;103(9):1642-56. Epub 2016 Sep 2.

Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 97331 USA.

Premise Of The Study: The basal asterid clade Cornales radiated during the Late Cretaceous. However, our understanding of early evolutionary patterns and relationships remain obscure. New data from five permineralized fruits in calcareous concretions from the Upper Cretaceous (Coniacian-Santonian) Haborogawa Formation, Hokkaido, Japan provide anatomical details that aid our knowledge of the group.

Methods: Specimens were studied from cellulose acetate peels, and three-dimensional reconstructions were rendered using AVIZO.

Key Results: Fruits are drupaceous, roughly pyriform, 2.9-4.3 mm in diameter, with a fleshy mesocarp, transition sclereids, and a stony endocarp of four to five locules, with the septa forming a cross or star-like pattern in transverse section, distinct germination valves, and one apically attached anatropous seed per locule. Vascular tissue occurs in zones between the mesocarp and exocarp, in two rows within the septa, and prominent seed bundles can be traced throughout the fruit sections. Seeds have a single integumentary layer of radially flattened square to rectangular cells and copious cellular endosperm. A fully formed, straight, cellular dicotyledonous embryo, with closely appressed, spathulate cotyledons, is present within each seed.

Conclusions: The unique combination of characters shown by these fruits is found in Cornaceae, Curtisiaceae, and Davidiaceae and allows us to describe a new taxon of Cornales, Eydeia hokkaidoensis gen. et sp. nov., with many similarities to extant Davidia involucrata. These fossils underscore the phylogenetic diversification of Cornales that was underway during the Late Cretaceous and support the hypothesis that a Davidia-like fruit morphology is plesiomorphic within Cornales.
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http://dx.doi.org/10.3732/ajb.1600151DOI Listing
September 2016

Phylogenetic diversification of Early Cretaceous seed plants: The compound seed cone of Doylea tetrahedrasperma.

Am J Bot 2016 05 11;103(5):923-37. Epub 2016 May 11.

Department of Botany and Plant Pathology, 2082 Cordley Hall, Oregon State University, Corvallis, Oregon 97331, USA.

Premise Of The Study: Discovery of cupulate ovules of Doylea tetrahedrasperma within a compact, compound seed cone highlights the rich diversity of fructification morphologies, pollination biologies, postpollination enclosure of seeds, and systematic diversity of Early Cretaceous gymnosperms.

Methods: Specimens were studied using the cellulose acetate peel technique, three-dimensional reconstructions (in AVIZO), and morphological phylogenetic analyses (in TNT).

Key Results: Doylea tetrahedrasperma has bract/fertile short shoot complexes helically arranged within a compact, compound seed cone. Complexes diverge from the axis as a single unit and separate distally into a free bract tip and two sporophylls. Each sporophyll bears a single, abaxial seed, recurved toward the cone axis, that is enveloped after pollinaton by sporophyll tissue, forming a closed cupule. Ovules are pollinated by bisaccate grains captured by micropylar pollination horns.

Conclusions: The unique combination of characters shown by D. tetrahedrasperma includes the presence of cupulate seeds borne in conifer-like compound seed cones, an ovuliferous scale analogue structurally equivalent to the ovulate stalk of Ginkgo biloba, gymnospermous pollination, and nearly complete enclosure of mature seeds. These features characterize the Doyleales ord. nov., clearly distinguish it from the seed fern order Corystospermales, and allow for recognition of another recently described Early Cretaceous seed plant as a second species in genus Doylea. A morphological phylogenetic analysis highlights systematic relationships of the Doyleales ord. nov. and emphasizes the explosive phylogenetic diversification of gymnosperms that was underway at the time when flowering plants may have originated and/or first began to radiate.
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http://dx.doi.org/10.3732/ajb.1600030DOI Listing
May 2016

Exploring the fossil history of pleurocarpous mosses: Tricostaceae fam. nov. from the Cretaceous of Vancouver Island, Canada.

Am J Bot 2015 Nov 5;102(11):1883-900. Epub 2015 Nov 5.

Department of Biological Sciences, Humboldt State University, Arcata, California 95221 USA

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).
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http://dx.doi.org/10.3732/ajb.1500360DOI Listing
November 2015

Hughmillerites vancouverensis sp. nov. and the Cretaceous diversification of Cupressaceae.

Am J Bot 2014 Dec 21;101(12):2136-47. Epub 2014 Nov 21.

Department of Botany and Plant Pathology, 2082 Cordley Hall, Oregon State University, Corvallis, Oregon 97331 USA.

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.
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http://dx.doi.org/10.3732/ajb.1400369DOI Listing
December 2014

Honeggeriella complexa gen. et sp. nov., a heteromerous lichen from the Lower Cretaceous of Vancouver Island (British Columbia, Canada).

Am J Bot 2013 Feb 10;100(2):450-9. Epub 2013 Jan 10.

Department of Biological Sciences, Humboldt State University, Arcata, CA 95521, USA.

Premise Of The Study: Colonists of even the most inhospitable environments, lichens are present in all terrestrial ecosystems. Because of their ecological versatility and ubiquity, they have been considered excellent candidates for early colonizers of terrestrial environments. Despite such predictions, good preservation potential, and the extant diversity of lichenized fungi, the fossil record of lichen associations is sparse. Unequivocal lichen fossils are rare due, in part, to difficulties in ascertaining the presence of both symbionts and in characterizing their interactions. This study describes an exceptionally well-preserved heteromerous lichen from the Lower Cretaceous of Vancouver Island.

Methods: The fossil occurs in a marine carbonate concretion collected from the Apple Bay locality on Vancouver Island, British Columbia, and was prepared for light microscopy and SEM using the cellulose acetate peel technique.

Key Results: The lichen, Honeggeriella complexa gen. et sp. nov., is formed by an ascomycete mycobiont and a chlorophyte photobiont, and exhibits heteromerous thallus organization. This is paired with a mycobiont-photobiont interface characterized by intracellular haustoria, previously not documented in the fossil record.

Conclusions: Honeggeriella adds a lichen component to one of the richest and best characterized Early Cretaceous floras and provides a significant addition to the sparse fossil record of lichens. As a heteromerous chlorolichen, it bridges the >350 million-year gap between previously documented Early Devonian and Eocene occurrences.
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http://dx.doi.org/10.3732/ajb.1200470DOI Listing
February 2013

Seed cone anatomy of Cheirolepidiaceae (Coniferales): reinterpreting Pararaucaria patagonica Wieland.

Am J Bot 2012 Jun 4;99(6):1058-68. Epub 2012 Jun 4.

CONICET-Museo Paleontologico Egidio Feruglio, Av. Fontana 140, Trelew, Chubut 9100, Argentina.

Premise Of The Study: Seed cone morphology and anatomy reflect some of the most important changes in the phylogeny and evolutionary biology of conifers. Reexamination of the enigmatic Jurassic seed cone Pararaucaria patagonica reveals previously unknown systematically informative characters that demonstrate affinities with the Cheirolepidiaceae. This paper documents, for the first time, internal anatomy for seed cones of this important extinct Mesozoic conifer family, which may represent the ghost lineage leading to modern Pinaceae.

Methods: Morphology and anatomy of cones from the Jurassic La Matilde Formation in Patagonia are described from a combination of polished wafers and thin section preparations. New photographic techniques are employed to reveal histological details of thin sections in which organic cell wall remains are not preserved. Specific terminology for conifer seed cones is proposed to help clarify hypotheses of homology for the various structures of the cones.

Key Results: Specimens are demonstrated to have trilobed ovuliferous scale tips along with a seed enclosing pocket of ovuliferous scale tissue. Originally thought to represent a seed wing in P. patagonica, this pocket-forming tissue is comparable to the flap of tissue covering seeds of compressed cheirolepidiaceous cones and is probably the most diagnostic character for seed cones of the family.

Conclusions: Pararaucaria patagonica is assigned to Cheirolepidiaceae, documenting anatomical features for seed cones of the family and providing evidence for the antiquity of pinoid conifers leading to the origin of Pinaceae. A list of key morphological and anatomical characters for seed cones of Cheirolepidiaceae is developed to facilitate assignment of a much broader range of fossil remains to the family. This confirms the presence of Cheirolepidiaceae in the Jurassic of the Southern Hemisphere, which was previously suspected from palynological records.
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http://dx.doi.org/10.3732/ajb.1100544DOI Listing
June 2012

A lower Cretaceous (Valanginian) seed cone provides the earliest fossil record for Picea (Pinaceae).

Am J Bot 2012 Jun 23;99(6):1069-82. Epub 2012 May 23.

Department of Biological Sciences, CW 405 Biological Sciences Building, University of Alberta, Edmonton, Alberta, Canada T6G 2E9.

Premise Of Study: Sequence analyses for Pinaceae have suggested that extant genera diverged in the late Mesozoic. While the fossil record indicates that Pinaceae was highly diverse during the Cretaceous, there are few records of living genera. This description of an anatomically preserved seed cone extends the fossil record for Picea A. Dietrich (Pinaceae) by ∼75 Ma.

Methods: The specimen was collected from the Apple Bay locality of Vancouver Island (Lower Cretaceous, Valanginian) and is described from anatomical sections prepared using cellulose acetate peels. Cladistic analyses of fossil and extant pinaceous seed cones employed parsimony ratchet searches of an anatomical and morphological matrix.

Key Results: This new seed cone has a combination of characters shared only with the genus Picea A. Dietr. and is thus described as Picea burtonii Klymiuk et Stockey sp. nov. Bisaccate pollen attributable to Picea is found in the micropyles of several ovules, corroborating the designation of this cone as an early spruce. Cladistic analyses place P. burtonii with extant Picea and an Oligocene representative of the genus. Furthermore, our analyses indicate that Picea is sister to Cathaya Chun et Kuang, and P. burtonii helps to establish a minimum date for this node in hypotheses of conifer phylogeny.

Conclusions: As an early member of the extant genus Picea, this seed cone extends the fossil record of Picea to the Valanginian Stage of the Early Cretaceous, ca. 136 Ma, thereby resolving a ghost lineage predicted by molecular divergence analyses, and offers new insight into the evolution of Pinaceae.
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http://dx.doi.org/10.3732/ajb.1100568DOI Listing
June 2012

The seed cone Eathiestrobus gen. nov.: fossil evidence for a Jurassic origin of Pinaceae.

Am J Bot 2012 Apr;99(4):708-20

Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701, USA.

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.
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http://dx.doi.org/10.3732/ajb.1100595DOI Listing
April 2012

A new family of leafy liverworts from the middle Eocene of Vancouver Island, British Columbia, Canada.

Am J Bot 2011 Jun 17;98(6):998-1006. Epub 2011 May 17.

Department of Biological Sciences, Humboldt State University, Arcata, California 95521, USA.

Premise Of The Study: Morphology is a reflection of evolution, and as the majority of biodiversity that has lived on Earth is now extinct, the study of the fossil record provides a more complete picture of evolution. This study investigates anatomically preserved bryophyte fossils from the Eocene Oyster Bay Formation of Vancouver Island. While the bryophyte fossil record is limited in general, anatomically preserved bryophytes are even more infrequent; thus, the Oyster Bay bryophytes are a particularly significant addition to the bryophyte fossil record.

Methods: Fossils occur in two marine carbonate nodules collected from the Appian Way locality on the eastern shore of Vancouver Island, British Columbia, and were prepared using the cellulose acetate peel technique.

Key Results: The fossils exhibit a novel combination of characters unknown among extinct and extant liverworts: (1) three-ranked helical phyllotaxis with underleaves larger than the lateral leaves; (2) fascicled rhizoids associated with the leaves of all three ranks; (3) Anomoclada-type endogenous branching.

Conclusions: A new liverwort family, Appianacae fam. nov., is established based upon the novel combination of characters. Appiana gen. nov. broadens the known diversity of bryophytes and adds a hepatic component to one of the richest and best characterized Eocene floras.
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http://dx.doi.org/10.3732/ajb.1000396DOI Listing
June 2011

Phylogenetic diversification of Equisetum (Equisetales) as inferred from Lower Cretaceous species of British Columbia, Canada.

Am J Bot 2009 Jul 12;96(7):1289-99. Epub 2009 Jun 12.

Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701 USA.

Three types of anatomically preserved vegetative shoots with features that characterize crown group Equisetum 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 Equisetum 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.
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http://dx.doi.org/10.3732/ajb.0800381DOI Listing
July 2009

Anatomy and development of fruits of Lauraceae from the Middle Eocene Princeton Chert.

Am J Bot 2009 Mar;96(3):637-51

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada.

Investigations of the Middle Eocene Princeton Chert reveal evidence for the connection of lauraceous flowers to fruits through a developmental series. Youngest fruits are found with attached floral remnants. Later stages show receptacle enlargement, fruit wall thickening, and the development of abundant sclereid clusters. Mature fruits are borne on a shallow receptacle and have an endocarp palisade layer of radially elongate cells with stellate outlines, an inner mesocarp layer of radiately arranged sclereid clusters, and a fleshy outer mesocarp layer containing numerous idioblasts with contents. Each mature fruit bears a single seed retaining the outer integument with an innermost radially elongate transfusion cell layer. Mature seeds contain a cellular embryo bearing idioblasts. Fruits are distinguishable from previously described anatomically preserved fossil taxa. This study represents the only documented developmental reconstruction of fossil fruits of Lauraceae and that self-pruning evolved prior to the Eocene. Anatomical modifications over the developmental sequence indicate that different stages of maturity preserved together, may be erroneously identified as several taxa at a fossil locality. Fossil morphotypes typically underestimate species number, but this study suggests that the number of inferred species based on fruit types may be inflated for Lauraceae, potentially exaggerating the tropical interpretation of the paleoenvironment.
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http://dx.doi.org/10.3732/ajb.0800318DOI Listing
March 2009

Distinguishing angiophytes from the earliest angiosperms: A Lower Cretaceous (Valanginian-Hauterivian) fruit-like reproductive structure.

Am J Bot 2009 Jan;96(1):323-35

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada T6G 2E9.

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 trichomes. They apparently broke open to release the tightly enclosed seeds by valves. Seeds are similar to those of the Triassic seed fern Petriellaea, but are about 100 million years younger and differ in size, vascularization, integumentary anatomy, seed attachment, and number of seeds/cupule. These new seeds are described as Doylea tetrahedrasperma 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.
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http://dx.doi.org/10.3732/ajb.0800295DOI Listing
January 2009

Is the anthophyte hypothesis alive and well? New evidence from the reproductive structures of Bennettitales.

Am J Bot 2009 Jan;96(1):296-322

Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701 USA.

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 carpel. Cladistic tests were performed to assess the strength of the "anthophyte hypothesis" and possible relationships 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.
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http://dx.doi.org/10.3732/ajb.0800209DOI Listing
January 2009

Introduction to the Darwin special issue: The abominable mystery1.

Am J Bot 2009 Jan;96(1):3-4

Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada.

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http://dx.doi.org/10.3732/ajb.0800402DOI Listing
January 2009

Todea from the Lower Cretaceous of western North America: implications for the phylogeny, systematics, and evolution of modern Osmundaceae.

Am J Bot 2008 Mar;95(3):330-9

Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701 USA.

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.
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http://dx.doi.org/10.3732/ajb.95.3.330DOI Listing
March 2008

Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov. (Saururaceae) from the Middle Eocene Princeton Chert.

Am J Bot 2007 Oct;94(10):1642-57

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada.

Investigations of small permineralized flowers from the Middle Eocene Princeton Chert, British Columbia, Canada have revealed that they represent an extinct species of Saururus. Over 100 flowers and one partial inflorescence were studied, and numerous minute perianthless flowers are borne in an indeterminate raceme. Each flower is subtended by a bract, and flowers and bracts are borne at the end of a common stalk. Five stamens are basally adnate to the carpels. Pollen is frequently found in situ in the anthers. Examined under SEM and TEM, pollen grains are minute (6-11 μm), monosulcate, boat-shaped-elliptic, with punctate sculpturing and a granulate aperture membrane. The gynoecium is composed of four basally connate, lobed carpels with recurved styles and a single ovule per carpel. Flower structure and pollen are indicative of Saururaceae (Piperales), and in phylogenetic analyses using morphological characters, the fossils are sister to extant Saururus. The fossil flowers are described here as Saururus tuckerae sp. nov. These fossil specimens add to the otherwise sparse fossil record of Piperales, represent the oldest fossils of Saururaceae as well as the first North American fossil specimens of this family, and provide the first evidence of saururaceous pollen in the fossil record.
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http://dx.doi.org/10.3732/ajb.94.10.1642DOI Listing
October 2007

Margaretbarromyces dictyosporus gen. sp. nov.: a permineralized corticolous ascomycete from the Eocene of Vancouver Island, British Columbia.

Mycol Res 2007 Jun 15;111(Pt 6):680-4. Epub 2007 Mar 15.

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada.

A single, permineralized ascoma resembling a pseudothecium assignable to the Pleosporales is described from the Eocene Appian Way fossil locality on Vancouver Island, British Columbia. The ascoma is globose, ostiolate, and erumpent on a fragment of the bark from an unidentified seed plant. Basally arranged asci contain large, multicelled, obovate ascospores within a single cavity or locule enclosed by a two-layered pseudoparenchymatous tissue that ostensibly represents ascostroma. Given this interpretation of the specimen's morphological features, Margaretbarromyces dictyosporus gen. sp. nov. represents the first report of a corticolous pleosporalean ascoma in the fossil record.
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http://dx.doi.org/10.1016/j.mycres.2007.03.010DOI Listing
June 2007

Cobbania corrugata gen. et comb. nov. (Araceae): a floating aquatic monocot from the Upper Cretaceous of western North America.

Am J Bot 2007 Apr;94(4):609-24

Department of Biological Sciences, University of Alberta, Edmonton, AB T6G 2E9 Canada;

The fossil record of aquatic flowering plants broadens our understanding of their former diversity and origins from terrestrial ancestors. This paper describes a floating aquatic monocot from 71 whole plants and several isolated leaf fragments from Upper Cretaceous oxbow lake sediments in the Dinosaur Park Formation, Alberta, Canada. The new material is represented by rosettes of leaves and roots attached to short stems that are interconnected by stolons and corresponds to the fossil aroid originally described as Pistia corrugata Lesquereux. Up to six plants have been found interconnected on a single slab suggesting that these plants grew in extensive floating mats covering lakes and calm stretches of rivers. Stems have up to six leaves and large numbers of branched aquatic roots. The leaf is trumpet-shaped with an elongate clasping petiole, large aerenchymatous base, and a nearly circular blade rim. Leaf bases are often filled with sediment giving the leaf the appearance of having a basal pouch. Petioles have 6-9 veins that divide into an upper and lower set, and veins converge at an apical notch. A submarginal collective vein and at least two marginal veins with branching veins form the leaf rim. A series of dichotomizing and anastomosing veins characterize the adaxial leaf surface. Tertiary and quaternary veins form polygonal areolae. Leaf surfaces are covered in trichomes that, like those in Pistia stratiotes, probably aided in buoyancy. A reconstruction of the plant is presented. Based on unique leaf morphology, these fossil plants are clearly not assignable to the genus Pistia and are described as Cobbania corrugata (Lesquereux) Stockey, Rothwell et Johnson gen. et comb. nov. Recent systematic analyses using molecular characters resolve two separate origins of floating aquatic aroids included in the duckweeds and the genus Pistia. This new fossil genus increases our understanding of colonization of aquatic habitats by revealing a third possible origin of the floating aquatic habit within Araceae.
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http://dx.doi.org/10.3732/ajb.94.4.609DOI Listing
April 2007

Cascadiacarpa spinosa gen. et sp. nov. (Fagaceae): castaneoid fruits from the Eocene of Vancouver Island, Canada.

Am J Bot 2007 Mar;94(3):351-61

Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada;

Documenting the paleodiversity of well-studied angiosperm families serves to broaden their circumscription while also providing a time-specific reference point to mark the first occurrence of characters and appearance of lineages. More than 80 anatomically preserved specimens of spiny, cupulate fruits in various developmental stages have been studied from the Eocene Appian Way locality of Vancouver Island, British Columbia, Canada. Details of internal anatomy and external morphology are known for the cupules, fruits, and pedicels. Cupule spines branch and are often borne in clusters. Cupules lack clear sutures and are adnate to a single nut that is enclosed entirely with the exception of the apical stylar protrusion of the pistil. A central hollow cylinder of vascular tissue can be seen extending up the peduncle to the base of the fruit and along the inner wall of the cupule. The fruit has a sclerotic outer pericarp that grades into a parenchymatous mesocarp and a sclerotic endocarp lining the locules. Early in development, the two locules are divided by a thin septum to which the ovules are attached. Only one seed develops to maturity as evidenced by an embryo occupying the locule alongside an abortive apical ovule. Three-dimensional reconstructions of these fruits have allowed for comparisons to both extinct and extant fagaceous taxa. The Appian Way fruits are most similar to extant Castanopsis species (Fagaceae) but differ in having only two locules. Cascadiacarpa spinosa gen. et sp. nov. Mindell, Stockey et Beard is the first occurence of a bipartite gynoecium and earliest known occurrence of hypogeous fruits in Fagaceae. The appearance of Casacadiacarpa in the Eocene of British Columbia supports a Paleogene radiation of the family. The numerous derived characters of these fruits show that evalvate, spiny, single-fruited cupules of Fagaceae were present in the Paleogene of North America.
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http://dx.doi.org/10.3732/ajb.94.3.351DOI Listing
March 2007

Solenostelopteris skogiae sp. nov. from the Lower Cretaceous of Vancouver Island.

J Plant Res 2006 Sep 29;119(5):525-32. Epub 2006 Aug 29.

Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada.

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.
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http://dx.doi.org/10.1007/s10265-006-0027-2DOI Listing
September 2006

Beardia vancouverensis gen. et sp. nov. (Juglandaceae): permineralized fruits from the Eocene of British Columbia.

Am J Bot 2006 Apr;93(4):557-65

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada.

Large numbers of permineralized juglandaceous fruits were identified in calcareous nodules from the Eocene Appian Way locality on Vancouver Island, British Columbia, Canada. The fruits, small dorsiventrally flattened nutlets, 4.5-7.0 mm long and 5.5-9.0 × 3-5 mm in diameter, were studied using cellulose acetate peels. They are wingless, ribbed, and have a lobed epicarp that surrounds the nutlet. Cells of the inner epicarp are thin-walled and traversed by a system of branching vascular strands. The stony nutlet wall is composed of fibers, with an outer layer of distinctive idioblasts. The fruits have a symmetry like that in Juglandaceae, subfamily Juglandoideae, tribe Platycaryeae, while the fibrous nut walls are like those of subfamily Engelhardioideae. This unique combination of characters indicates that these fruits represent a new genus and species of Juglandaceae: Beardia vancouverensis gen. et sp. nov. The excellent preservation of the Appian Way specimens has allowed a unique view of the internal fruit anatomy and external morphology. As the only wingless, flattened nuts known in the family, they further extend the range of morphological variation in fruits in the family. These fossils further support the hypothesis that North America was an important center of generic diversity for Juglandaceae during the early Tertiary.
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http://dx.doi.org/10.3732/ajb.93.4.557DOI Listing
April 2006