Ruth Stockey
Oregon State University, Botany and Plant Pathology, Faculty Member
Research Interests: Botany, Evolutionary Biology, Plant Biology, Biology, Pollen, and 3 morePlant sciences, Cupressaceae, and Twig
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PremiseAn 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... more
PremiseAn 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.MethodsThe specimen was studied from serial transverse sections prepared by the cellulose acetate peel technique.ResultsThe 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, the...
Research Interests: Botany, Evolutionary Biology, Plant Biology, Biology, Ecology, and 5 moreCretaceous, Xylem, fERN, Rhizome, and Stele
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... more
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.
Research Interests: Evolutionary Biology, Paleontology, Plant Biology, Biology, Ecology, and 4 moreMedicine, Cretaceous, Arctic, and Moss
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Research Interests: Phylogenetics and Biology
Research Interests: Geology and Paleontology
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Research Interests: Geology and Cretaceous
PREMISE OF THE STUDYFossils show that fly-speck fungi have been reproducing with small, black thyriothecia on leaf surfaces for ∼250 million years. We analyze morphological characters of extant thyriothecial fungi to develop a... more
PREMISE OF THE STUDYFossils show that fly-speck fungi have been reproducing with small, black thyriothecia on leaf surfaces for ∼250 million years. We analyze morphological characters of extant thyriothecial fungi to develop a phylogenetic framework for interpreting fossil taxa.METHODSWe placed 59 extant fly-speck fungi in a phylogeny of 320 Ascomycota using nuclear ribosomal large and small subunit sequences, including newly determined sequences from nine taxa. We reconstructed ancestral character states using BayesTraits and maximum likelihood after coding 11 morphological characters based on original observations and literature. We analyzed the relationships of three previously published Mesozoic fossils using parsimony and our morphological character matrix, constrained by the molecular phylogeny.KEY RESULTSThyriothecia evolved convergently in multiple lineages of superficial, leaf- inhabiting ascomycetes. The radiate and ostiolate scutellum organization is restricted to Dothide...
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Premise of research. Four small pollen cones bearing pollen have been found attached to a leafy twig in a calcareous concretion from the Valanginian Apple Bay locality, northern Vancouver Island, British Columbia, increasing the known... more
Premise of research. Four small pollen cones bearing pollen have been found attached to a leafy twig in a calcareous concretion from the Valanginian Apple Bay locality, northern Vancouver Island, British Columbia, increasing the known morphological diversity of Early Cretaceous cupressaceous pollen cones. Methodology. Specimens were prepared using the cellulose acetate peel technique, pollen was examined using SEM, and three-dimensional reconstructions were rendered with Avizo. Pivotal results. The small, subglobose cones bear helically arranged microsporophylls with a central resin canal. Each sporophyll bears three abaxial pollen sacs. Walls of pollen sacs are comprised of radially elongated rectangular cells with prominent wall thickenings. Most pollen sacs are intact and contain numerous, nonsaccate pollen grains. Pollen is 16–24 μm in diameter with one exit papilla that forms a low protuberance on the grain. External surfaces of grains are scabrate with numerous orbicules and macrogranules, and the nexine is laminated. Cone and pollen morphology most closely resemble taxa of the taxodioid Cupressaceae. The number of pollen sacs per microsporophyll is similar to Athrotaxis, Metasequoia, Sequoia, Sequoiadendron, and the Paleogene fossil conifer Homalcoia. Arrangement of cones on branches differs from those of Athrotaxis, Metasequoia, and Homalcoia; pollen lacks the long distal papilla of Sequoia and Sequoiadendron and is most similar in size to that in some cupressoid Cupressaceae in Chamaecyparis, Thuja, and Juniperus but with a smaller aperture. Conclusions. This combination of features is unlike that found in any living or extinct species of Cupressaceae. The fossil species is described as Morinostrobus holbergensis gen. et sp. nov., furthering our understanding of character evolution within Cupressaceae s.l. and adding to the evidence that the Lower Cretaceous was a time of increasing cupressaceous diversity.
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Premise of research. A large number of floating aquatic aroid fossils have been recovered from pond sediments in the Hell Creek Formation (Upper Cretaceous) of South Dakota, providing valuable new data about aquatic vegetation of the... more
Premise of research. A large number of floating aquatic aroid fossils have been recovered from pond sediments in the Hell Creek Formation (Upper Cretaceous) of South Dakota, providing valuable new data about aquatic vegetation of the uppermost Cretaceous, that are used to describe a new species of the genus Cobbania, and to evaluate associated reproductive structures and phylogenetic relationships among floating aquatic monocots. Methodology. Fossils were uncovered as needed with fine needles to reveal surface features of the specimens. Images were captured with a digital scanning camera, and phylogenetic analyses were conducted with TNT implemented through WinClada. Pivotal results. The new species, Cobbania hickeyi Stockey, Rothwell & Johnson, extends the range of the genus to the uppermost Cretaceous, supports the taxonomic integrity of the genus Cobbania, and increases our understanding of structural variation and species richness within the genus. Associated reproductive structures include an aroid spadix, strengthening the assignment of Cobbania to the Araceae. Phylogenetic analyses using “total-evidence” data help resolve conflicting results from either morphological or nucleotide sequence analyses of relationships among floating aquatic aroids, and the fossil taxon Aquaephyllum does not nest among the other floating aquatic species in any of our results. Conclusions. Species of the genus Cobbania were an important component of aquatic vegetation across the Northern Hemisphere during the Late Cretaceous. In tests of competing hypotheses for relationships among Pistia stratiotes, Cobbania spp., and species of Araceae subfamily Lemnoideae, the results from a “total-evidence” analysis suggest that specializations for the floating aquatic life form may be overwhelming other characters in the results of morphological analyses alone.
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Fossil angiosperm wood was collected from shallow marine deposits in the Upper Cretaceous (Coniacian) Comox Formation on Vancouver Island, British Columbia, Canada. The largest specimen is a log at least 2 m long and 38 cm in diameter.... more
Fossil angiosperm wood was collected from shallow marine deposits in the Upper Cretaceous (Coniacian) Comox Formation on Vancouver Island, British Columbia, Canada. The largest specimen is a log at least 2 m long and 38 cm in diameter. Thin sections from a sample of this log reveal diffuseporous wood with indistinct growth rings and anatomy similar toParaphyllanthoxylon. Occasional idioblasts with dark contents in the rays distinguish this wood from previously knownParaphyllanthoxylonspecies and suggest affinity with Lauraceae. The log also includes galleries filled with dry-wood termite coprolites. This trunk reveals the presence of tree-sized angiosperms in what is now British Columbia, and the association of dry-wood termites with angiosperm woods by the Coniacian (89 Ma). To understand the significance of this discovery, we reviewed the record of Cretaceous woods from North America. Our analysis of the distribution of fossil wood occurrences from Cretaceous deposits supports the...
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Premise of research. Two permineralized fruits have been found within a single calcareous concretion from the Upper Cretaceous (upper Campanian) Spray Formation of Vancouver Island, British Columbia. These fossils provide additional... more
Premise of research. Two permineralized fruits have been found within a single calcareous concretion from the Upper Cretaceous (upper Campanian) Spray Formation of Vancouver Island, British Columbia. These fossils provide additional evidence for the initial phylogenetic diversification of Cornales, the earliest diverging asterid lineage. Methodology. Specimens were prepared using the cellulose acetate peel technique. Pivotal results. A new cornalean species, Suciacarpa xiangae sp. nov., is characterized on the basis of a well-preserved fruit specimen. This fruit is represented by a large ovoid tetralocular sclerenchymatous endocarp. Locules are crescent shaped in transverse section and accompanied by short dorsal germination valves. Valves consist of small isodiametric sclereids as well as a large number of large secretory cavities. The septum and central axis are composed of transversely oriented and interwoven fibers and elongate sclereids. There is no major central vascular bundle; however, there are numerous small bundles scattered within the septa. This fruit has one apically attached seed per locule. An additional cornalean taxon—Sheltercarpa vancouverensis gen. et sp. nov., also represented by a single smooth tetralocular sclerenchymatous endocarp—consists of isodiametric and a few elongate sclereids, with short germination valves, one seed per locule, and lacks a central vascular bundle. Conclusions. Because of unique combinations of characters, the two fruits described in this study represent two new cornalean taxa, S. xiangae sp. nov. and S. vancouverensis gen. et sp. nov. These fruits display mosaics of apomorphic and plesiomorphic characters that occur in families of at least two different major cornalean clades: Cornaceae/Alangiaceae and Nyssaceae/Mastixiaceae/Davidiaceae. The morphological diversity seen in these taxa and other cornaleans from the Late Cretaceous suggests that the Campanian was an important time for the initial phylogenetic diversification of Cornales.
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Relationships of living and fossil Pinaceae were inferred using parsimony and Bayesian inference of morphological characters and plastid and nuclear DNA sequences. When considering extant taxa only, adding molecular to morphological... more
Relationships of living and fossil Pinaceae were inferred using parsimony and Bayesian inference of morphological characters and plastid and nuclear DNA sequences. When considering extant taxa only, adding molecular to morphological characters resulted in markedly increased resolution and branch support compared with analysis of morphology alone. Including 45 fossil taxa resulted in drastically decreased resolution in morphology-based consensus trees. We evaluated the effect on branch support and resolution of including DNA sequences, deleting fossils lacking information for cone scale apices and seeds, using reduced consensus methods, and using implied weighting, and found that the greatest improvements were found by including DNA sequences and using implied weighting. The tree topologies from parsimony and Bayesian inference confirm previous findings that the fossil genus Pseudoaraucaria and a few species of Pityostrobus from the Lower Cretaceous are related to abietoid genera, an...
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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... more
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. Specimens were studied from cellulose acetate peels, and three-dimensional reconstructions were rendered using AVIZO. 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 sing...
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The evolutionary history of Pinaceae is documented by numerous fossils, which include a number of anatomically preserved ovulate cones with many systematically informative characters. To date, four extinct genera have been recognized:... more
The evolutionary history of Pinaceae is documented by numerous fossils, which include a number of anatomically preserved ovulate cones with many systematically informative characters. To date, four extinct genera have been recognized: Pseudoaraucaria, Pityostrobus, Obirastrobus and Eathiestrobus. A new pinaceous cone from the Early Cretaceous of California is described as Pityostrobus pluriresinosa sp. nov., adding to the diversity of known Cretaceous pinaceous cones. Phylogenetic analysis places the new fossil cone in the pinoid clade along with Pinus, Eathiestrobus and numerous species of Pityostrobus. Combined with a non-metric multidimensional scaling ordination, the data reinforce that while Pityostrobus does not represent a natural genus, it likely reflects the rapid evolution of Pinaceae by showing that ovulate cone morphospace was widely occupied by Pityostrobus that has become more restricted in extant genera. Early Cretaceous taxa from Europe and eastern North America show the broadest occupation of morphospace, which then became more restricted to morphologies typical of the pinoid clade in the Late Cretaceous. Taxa from western North America and Asia are more similar to extant genera and occupy a smaller part of the morphospace, highlighting that eastern North America–Europe probably represents an original centre of early diversification for Pinaceae.
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Several anatomieally preserved twigs, a branehing speeimen and the wood of a large axis with affinities to Rosaeeae are deseribed from the Prineeton ehert (Middle Eoeene) of British Columbia, Canada. Speeimens are eharaeterised by a... more
Several anatomieally preserved twigs, a branehing speeimen and the wood of a large axis with affinities to Rosaeeae are deseribed from the Prineeton ehert (Middle Eoeene) of British Columbia, Canada. Speeimens are eharaeterised by a heteroeellular pith with a peri-medullary rone of thiek-walled oval eells and semi-ring-porous seeondary xylem with vertieal traumatie duets, fibres with eireular bordered pits, and mostly seanty paratracheal and oeeasionally apotracheal parenehyma. Ray to vessel pitting is similar to the alternate intervaseular pitting. Seeondary phloem is eomposed of tangentially oriented diseontinuous bands of alternating fibres and thinwalled eells. Seeondary eortical tissues are represented by a phelloderm eharaeterised by rectangular eells and phellern with rectangular eoneave eells. Anatomical variation between speeimens can be related to age of the woody axes. Juvenile and mature wood of this speeies differ in vessel arrangement and presenee of scanty paratrachea...
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Research Interests: Botany, Evolutionary Biology, Plant Biology, Biology, Cretaceous, and 4 morePlant sciences, Moss, Bryophyte, and Bay
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Page 1. BOT. GAZ. 147(4):508-548. 1986. C 1986 by The University of Chicago. All rights reserved. 0006-8071/86/4704-0004$02.00 CUTICLE MICROMORPH OLOGY OF ARAUCARIA DE JUSSIEU RUTH A. STOCKEY AND ...
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An ovulate pinaceous cone and associated needles, wood, and pollen cones were found permineralized within a small lens of chert in sediments of Middle Eocene age near Smithers, B.C. The ovulate cone, Pinus driftwoodensis sp.n., is... more
An ovulate pinaceous cone and associated needles, wood, and pollen cones were found permineralized within a small lens of chert in sediments of Middle Eocene age near Smithers, B.C. The ovulate cone, Pinus driftwoodensis sp.n., is cylindrical, about 3 cm long and 2.7 cm in diameter. Ovuliferous scales have swollen apices and an umbo lacking a spine. Pith and cortex of the axis are parenchymatous. The narrow vascular cylinder includes a ring of up to 70 resin canals. The trace to each bract and its associated ovuliferous scale arises as a single unit, circular in cross section, with an abaxial resin canal system. Two ovules with well-developed integuments but little internal tissue are borne on each ovuliferous scale. Pinaceous leaves, in two-needle fascicles, are associated with the cone. These needles contain two vascular bundles, numerous medial resin canals, stomata on both abaxial and adaxial surfaces, and a uniform hypodermis. Associated wood and small twigs have pinoid resin canals, heterogeneous rays to 14 cells high, uniseriate circular-bordered pits, and pinoid pits in the cross field, and appear similar to that described for P. similkarneenensis Miller. Pollen cones are 6 mm long and cylindrical and have dehisced prior to preservation. The ovulate cone and the two-needle fascicles are closely comparable to members of the genus Pinus, subgenus Pinus, section Pinus, subsections Sylvestres, Ponderosae, or Australes and may represent parts of the same plant.
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Several hundred vegetative and fertile specimens of Azolla Lam. have been recovered from the Paleocene Paskapoo Formation at the Joffre Bridge locality (Middle Tiffanian (Ti3) age) near Red Deer, Alberta. The spore complexes closely... more
Several hundred vegetative and fertile specimens of Azolla Lam. have been recovered from the Paleocene Paskapoo Formation at the Joffre Bridge locality (Middle Tiffanian (Ti3) age) near Red Deer, Alberta. The spore complexes closely resemble those of the Paleocene A. stanleyi Jain & Hall, and the vegetative material is referred to that species. The specimens are unusually complete in that the remains of the fragile sporophyte are preserved, commonly with reproductive structures in place. Plants reaching up to 2.25 cm in length consist of alternately branched rhizomes bearing alternate, imbricate, sessile leaves. Leaves are ovate with entire margins, papillate surfaces, and a single midvein. Reproductive structures have been examined using light, scanning, and transmission electron microscopy. This new material is compared with the other Paleocene species for which sporophytes are known and discussed in terms of evolutionary trends for the genus. The specimens suggest that most of th...